Iv ... MYCOTAXON 136(4)

MYCOTAXON

VOLUME ONE HUNDRED THIRTY-SIX (4) — TABLE OF CONTENTS

SOPMCC HAG RD Fs. sons_2 uy tes tas cate chee et Rp 9 Go 9 a Yo skein aol sede, 2 eo Rees ae vi

Nomenclatural noveltiese? typifications sco. «08s a foc bee shoe 5A Dee lee vii

PROVIEW ETS at Soe cen cache 0 nfo ce age Wen een tone sat hae REE Y Hee Wed aha Hemiigh toon ae ren lang ix

DODD SOMESSTON PHOCOAULE 5. oi 52 a8 en voles teed Marat aded oie neu dees as et x

PROUT Te=FditOR <5 eh Res nth ei ea se Re Beats Aca 5 Sues oie a eee Ren Be xi

In Remembrance

Grégoire HenneberrO 29-2021) inc. hig kte Son mares xiii

JACK ROGEPS OS 7 S202) sy acn saha.g seeks le ehh eae thee Be XXXi

Kalida: Vaitky, 930-2021) cs cua eve veto ese eaen ean XXXiX

NEW SPECIES

Lamprospora benkertii sp. nov., and an evaluation of Lamprospora spp.

with seaveri-type ascospore ornamentation

JAN ECKSTEIN, MARCEL VEGA, ZUZANA SOCHOROVA, LUKAS JANOSIK 693

Pertusaria wui sp. nov. on bamboo from Yunnan, China

JIARONG ZHANG, LIN Liu, XIANDONG XUE, QIANG REN 719

Clavaria cystidiata sp. nov. from India

K. KRISHNAPRIYA & T.K. ARUN KUMAR 725

Aspicilia lixianensis, A. nivalis, and A. pycnocarpa spp. nov. from China

LIN LIU & QIANG REN 739

Architrypethelium barrerae sp. nov.

from a cloud forest in Veracruz, Mexico

JORGE GUZMAN-GUILLERMO & REGULO CARLOS LLARENA HERNANDEZ 749

Flavodontia rosea gen. & sp. nov. from southwestern China

Hui WANG & CHANG-LIN ZHAO 755

Pseudodeightoniella indica gen. and sp. nov.,

a hyphomycete from India SANJEET KUMAR VERMA, SANJAY YADAV,

RAGHVENDRA SINGH, BALMUKUND CHAURASIA, SHAMBHU KUMAR 769

Lepra taiwanensis and Lepra yunlingensis spp. nov. from China

JIARONG ZHANG, XIANDONG XUE, LIN Liu, XUYUN QIU, QIANG REN 779

VALIDATIONS

Validation of five Peronospora species names

WILLIAM J. Davis & JO ANNE CROUCH 785

OCTOBER-DECEMBER 2021 ... V

TYPIFICATIONS

Typification of Agaricus cespitosus, Ag. oniscus, and Ag. sphagnicola and their

synonymy with Lichenomphalia umbellifera ANDRUS VOITK 789

NEW RANGES/HOSTS

New records from Turkey: Cortinarius barbatus, C. osmophorus,

C. saturninus IsMAIL ACAR 819

Notes on lichenized fungi of chroodiscoid Thelotremataceae

from China ZE-FENG J1A, MIN Li, Yu-Ru Fu, Jinc Pu 831

Additions to the knowledge of foliicolous micromycetes in Turkey

GOKHAN DOGAN, MAKBULE ERDOGDU,

ZEKI AYTAC, ALI IHSAN KARAYEL, ZEKIYE SULUDERE 841

Cribraria lepida, Physarum dictyosporum, P. diderma, and P. spectabile

newly recorded from Turkey GONUL EROGLU 853

First report of Podosphaera cercidiphylli on endangered

Cercidiphyllum japonicum in China

Z1-JIA PENG, SHU-YAN Liu, ZHONG-Donc Yu, MEI Qt 865

MycosBioTa (FUNGAE) NEW TO THE MYCOTAXON WEBSITE

Macromycetes from woodland zones of

Milpa Alta mayoralty, Mexico City, Mexico (suMMaRy)

SIGFRIDO SIERRA, SANDRA CASTRO-SANTIUSTE,

IBETH RODRIGUEZ-GUTIERREZ, ARELI E. GONZALEZ-MENDOZA,

Mario AARON GUTIERREZ-SANCHEZ, LISETTE CHAVEZ-GARCIA,

DANIELA ABIGAIL GUZMAN-RAMIREZ, JOSE DE JESUS RuIZ-RAMOS,

GUADALUPE GALVAN-BECERRIL, NAVITH ALEJANDRA LOPEZ-GARDUZA,

LILIA PEREZ-RAMIREZ, JOAQUIN CIFUENTES 875

A checklist of Agaricus from Pakistan (summary)

ABDUL REHMAN NIAZI, ANEEQA GHAFOOR, HIRA BASHIR 877

BOOK REVIEWS AND NOTICES ELsE C. VELLINGA 879

VI ... MYCOTAXON 136(4)

CORRIGENDA

MYCOTAXON 136(2)

p.344, line 27 —— For: Collybiopsis subnuda (Ellis ex Peck) R.H. Petersen, comb. nov

IF 556198

Basionym: Marasmius subnudus Ellis ex Peck, 1898.

Bull. Torrey Bot. Club 25: 287.

[An invalid combination due to incorrect basionym citation]

READ: Collybiopsis subnuda (Ellis ex Peck) R.H. Petersen, comb. nov.

IF 556112

Basionym: Marasmius subnudus Ellis ex Peck, 1898.

Ann. Rep. New York St. Mus. 51: 287. 1898.

Validated on 6 February 2022 in INDEx FUNGORUM no. 510

http://www.indexfungorum.org/Publications/Index%20Fungorum%20no0.510.pdf

PUBLICATION DATE FOR VOLUME ONE HUNDRED THIRTY-SIX (3)

MYCOTAXON for JULY-SEPTEMBER 2020 (I-xII + 545-692)

was issued on November 4, 2021

OCTOBER-DECEMBER 2021 ...

NOMENCLATURAL NOVELTIES AND TYPIFICATIONS

PROPOSED IN MYCOTAXON 136(4)

Agaricus cespitosus Bolton 1836 (lectotypified),

[MBT 10004616], p. 792

Agaricus oniscus Fr. 1818 (lectotypified),

[MBT 10004612], p. 802 & (epitypified)

[MBT 10004614], p.802

Agaricus sphagnicola Berk. 1836 (neotypified),

[MBT 10004615], p. 814

Architrypethelium barrerae Guzm.-Guill. & Llar.-Hern.

[MB 837958], p. 750

Aspicilia lixianensis Q. Ren & Lin Liu

[FN 570763], p. 744

Aspicilia nivalis Q. Ren & Lin Liu

[FN 570764], p. 745

Aspicilia pycnocarpa Q. Ren & Lin Liu

[FN 570766], p. 742

Clavaria cystidiata Krishnapriya & T.K.A. Kumar

[MB 832680], p 728

Flavodontia C.L. Zhao

[MB 838322], p. 760

Flavodontia rosea C.L. Zhao

[MB 838323], p. 762

Lamprospora benkertii Eckstein, M. Vega, Sochorova & Janosik

[MB 831135], p. 696

Lepra taiwanensis Q. Ren

[EN 570770], p. 782

Lepra yunlingensis Q. Ren

[FN 570769], p. 780

Peronospora coronillae-minimae Vienn.-Bourg. ex W.J. Davis & J.A. Crouch

[MB 838139], p. 786

“Peronospora coronillae-minimae” Vienn.-Bourg., nom. inval.

Peronospora erini Vienn.-Bourg. ex W.J. Davis & J.A. Crouch

[MB 838140], p. 786

“Peronospora erinii’ Vienn.-Bourg., nom. inval.

VII

vill ... MYCOTAXON 136(4)

Peronospora papaveris-pilosi Vienn.-Bourg. ex W.J. Davis & J.A. Crouch

[MB 838141], p. 787

”Peronospora papaveris-pilosi” Vienn.-Bourg., nom. inval.

Peronospora ranunculi-flabellati Vienn.-Bourg. ex W.J. Davis & J.A. Crouch

[MB 838142], p. 787

” Peronospora ranunculi-flabellati” Vienn.-Bourg., nom. inval.

Peronospora viciicola L. Camp. ex W.J. Davis & J.A. Crouch

[MB 838143], p. 787

”Peronospora viciicola” L. Camp. [as ‘vicicola’] nom. inval.

Pertusaria wui Q. Ren

[FN 570761], p. 720

Pseudodeightoniella S.K. Verma, Sanj. Yadav & Raghv. Singh

[MB 836237], p. 772

Pseudodeightoniella indica S.K. Verma, Sanj. Yadav & Raghv. Singh

[MB 836238], p. 772

OCTOBER-DECEMBER 2021...

REVIEWERS — VOLUME ONE HUNDRED THIRTY-SIX (4)

The Editors express their appreciation to the following individuals who have,

prior to acceptance for publication, reviewed one or more of the papers

prepared for this issue.

Sabeena Aliyarukunju

Sinan Alkan

Andre Aptroot

Lu-chao Bai

Jordan Bailey

Curtis R. Bjork

Alisson Cardoso Rodrigues da Cruz

Balint Dima

Peter Dobbeler

Patricia Oliveira Fiuza

Mei-Ling Han

Man-Rong Huang

Sana Jabeen

Zefeng Jia

Ivona Kautmanova

Abdullah Kaya

Alejandro Kong Luz

Huajie Liu

Xiao-yong Liu

Juan Luis Mata

Renato Lucio Mendes Alvarenga

Pierre-Arthur Moreau

Lorelei L. Norvell

Shaun R. Pennycook

Meike Piepenbring

Luis A. Ramirez-Camejo

Qiang Ren

Amy Y. Rossman

Anton Shiryaev

Nina Shishkoff

Steve Stephenson

R. Greg Thorn

Nicolas Van Vooren

Lulu Zhang

Xin Zhao

x ... MYCOTAXON 136(4)

2022 MyCOTAXON SUBMISSION PROCEDURE

Prospective MycotTaxon authors should download the MycoTaxon 2022 guide,

review & submission forms, and MycoTaxon sample manuscript by clicking the ‘file

download page’ link on our INSTRUCTIONS TO AUTHORS page before preparing their

manuscript. This page briefly summarizes our “4-step’ submission process.

1—PEER REVIEW: Authors first contact peer reviewers (two for journal papers;

three for mycobiota/fungae) before sending them formatted text & illustration

files and the appropriate 2022 MycoTaxon journal or mycota reviewer comment

form. Experts return revisions & comments to BoTH the Editor-in-Chief

<editor@mycotaxon.com> and authors. ALL co-authors Must correct and proof-

read their files before submitting them to the Nomenclature Editor.

2—NOMENCLATURAL REVIEW: Authors email all ERROR-FREE text & illustration

files to the Nomenclature Editor <PennycookS@LandcareResearch.co.nz>.

Place first author surname + genus + ‘Mycotaxon’ on the subject line, and

(required) attach a completed sUBMISSION FORM. The Nomenclature Editor will

(i) immediately assign the accession number and (ii) after a few weeks return his

notes and suggested revisions to the author(s) and Editor-in-Chief.

3—FINAL SUBMISSION: All coauthors thoroughly revise and proof-read files

to prepare error-free text and images ready for immediate publication. Poorly

formatted copy willbe rejected or returned for revision. E-mail the final manuscript

to the Editor-in-Chief <editor@mycotaxon.com>, adding the accession number to

the message and all files, which include a (i) revised 2022 submission form, all (ii)

text files and (iii) jpg images, and (iv) FN, IF, or MB identifier verifications for each

new name or typification. The Editor-in-Chief acknowledges submissions within

two weeks of final submission but requests authors to wait at least 14 days before

sending a follow-up query (without attachments).

4—FINAL EDITORIAL REVIEW & PUBLICATION: The Editor-in-Chief conducts a

final grammatical and scientific review and returns her editorial revisions to all

expert reviewers and coauthors for final author approval. Author-approved files

are placed in the publication queue.

The PDF proof and bibliographic & nomenclatural index entries are sent to all

coauthors for final inspection. After PDF processing, the Editor-in-Chief corrects

ONLY PDF editorial/conversion and index entry errors; corrections of all other

errors are listed in the ERRATA of a subsequent issue for no charge. Authors will pay

fees for mycobiota uploads, optional open access, and correction of major author

errors to the Business Manager <subscriptions@mycotaxon.com> at this time.

MyYcOTAXON LTD— www.mycotaxon.com

The Mycotaxon Webmaster <mycotaxon@gmail.com> posts announcements,

subscription & publication information, and author forms & templates on the official

MycoTAXxon site. Our server also hosts the mycobiota web-page for free download

of Fungae (regional annotated species lists).

MYCOTAXON ONLINE— www.ingentaconnect.com/content/mtax/mt

The MycoTaxon journal publishes four quarterly issues per year. Both open access

and subscription articles are offered.

OCTOBER-DECEMBER 2021 ... XI

FROM THE EDITOR-IN-CHIEF

THREE MYCOLOGICAL GIANTS HAVE THIS WAY PASSED—Our remembrance

section covers the lives of three men who made outstanding contributions

both to mycology and to this journal. During 2021 we lost on June 14

Jack Davip RoGers (age 83), beloved dean of Xylariaceae and 1990-1998

Mycotaxon Editorial Advisory Board member; on October 12 KALMAN

GEzA VANKY (age 91), the world’s leading authority on the smut fungi

(Ustilaginomycotina) who published 56 significant papers in MyCOTAXON;

and on October 13 GREGOIRE LAURENT HENNEBERT (age 92), who with

Dick Korf founded Mycotaxon in 1974 and served as Book Review

Editor (1974-1991) and French Language Editor (1974-2006). Assembling

the photographs and researching the background to add to personal

reminiscences extended our already seriously late October-December 2021

issue, but we could not permit their histories to go uncelebrated or postponed.

DELETING THOSE PESKY PRIME SYMBOLS—For several years, our authors

and your editors have struggled over properly formatted latitude/longitude

coordinates. The process has been particularly onerous on the editorial side

because primes visible in MSWord Times/Times New Roman transform

into peculiar, seemingly unrelated symbols in the Editor-in-Chief’s Adobe

InDesign® program. Until recently, global replacements could sometimes

handle corrections in the program, but an unfortunate recent Adobe® update

now transforms ALL single/double prime symbols as empty pink boxes,

requiring tedious individual replacement of each symbol one by one—no

doubt portending Adobe’s intended removal of the ‘symbols’ fonts, certain to

make future formatting even more dire.

Therefore, MYCOTAXON issues a NEW REQUIREMENT for lat/long coordinates

that should reduce everyone's agony by eliminating the old degree/minute/

second presentation style. Henceforth:

(1) Authors should present their data as degrees with fractions rounded

off to four decimal places. [NoTE: 1° = <111 km, and 0.0001° = <11 m,

so any additional decimal places represent spurious accuracy. We shall

definitely accept coordinates rounded to fewer than four decimal places. ]

(2) No space should be inserted between the data and its cardinal letter

[N, E, S, W].

(3) Place one space but No punctuation between the latitude and longitude

components of a single set of coordinates. Example:

OLp and/or INCORRECT: 38°10°46"N, 13°15 ’40°E

NEw: 38.1794°N 13.2611°E.

XII ... MYCOTAXON 136(4)

THE 2021 OCTOBER-DECEMBER MYCoTAXON Offers 18 contributions by 60

authors (from 9 countries) revised by 35 expert reviewers and the editors.

The NEW TAXA section proposes Two new genera (Flavodontia from

China & Pseudodeightoniella from India) and ELEVEN species new to science

representing Architrypethelium from Mexico; Aspicilia, Flavodontia, Lepra,

and Pertusaria from CuHtNna; and Clavaria and Pseudodeightoniella from

INDIA.

VALIDATION presents a single paper validating five Peronospora species.

TYPIFICATIONS, also comprising one paper (but a long one), sets forth an

entertainingly convincing history that establishes three Fries, Bolton, and

Berkeley Agaricus names as synonyms of the lovely and nomenclaturally

controversial Lichenomphalia umbellifera.

NEW RANGES/HOSTS reports range extensions for [ascomycetes] Ascochyta,

Mycosphaerella, Phyllosticta, and Rhabdospora in TurKEy and Podosphaera in

Curina; [basidiomycetes] Cortinarius in TurKEy; [lichens] Astrochapsa and

Chapsa in Cuina; and [myxomycetes] Cribraria and Physarum in TURKEY.

Our issue concludes with the announcement of two new mycobiota for the

MycoTaxon website that cover macrofungi in Mexico City and Agaricus

species in Pakistan.

MycoTaxon 136(4) also provides identification keys to Pertusaria and

Clavaria species. Papers providing conclusions supported by sequence

analyses cover the new Aspicilia, Clavaria, Flavodontia, and Lamprospora

species and the new records for Cortinarius in Turkey and Podosphaera in

China.

Warm regards,

Lorelei L. Norvell (Editor-in-Chief)

6 February 2022

OCTOBER-DECEMBER 2021 ... XIII

IN CELEBRATION

MYCOTAXON BIDS AN AFFECTIONATE

FAREWELL TO OUR LAST CO-FOUNDER:

It is with profound sadness we note

the loss of GREGOIRE LAURENT |

HENNEBERT on 13 October 2021 at :

theage of 92. Weshall missthis gentle 2

soul, who was the consummate E

mycologist’s mycologist, master g

of both fungal taxonomy and 3

nomenclature, tireless collector,

outstanding fungal ambassador, and Fic. 1. Professor Hennebert at age 50 in 1979 on the

brilliant scholar. ‘new campus in Louvain-la-Neuve.

Gregoire’s early work was sparked by Prof. Victor Estienne, who commented

that scientists recognized several different Botrytis species on the onion

(Allium cepa), but “What are they, exactly? Are they not all Botrytis cinerea?”

His advisor recommended that the young Hennebert research host specificity

by comparing the different ‘taxa through cross-inoculations in host plants.”

A good suggestion, perhaps, except that these “cross-inoculations failed.”

Aconversation with Niels Buchwald (1949: SrUDIES OF SCLEROTINIACEAE)

turned Grégoire’s attention toward a different direction. “The matter is

primarily taxonomical. There are many names of Botrytis species in the

literature. Many are not Botrytis, some are, and among these, a number

are host specific, others are not. Differentiation of the species is first of all

MORPHOLOGICAL. You should compare fresh collections morphologically,

from nature and in culture. You should look for the perfect stage in nature

and find out the correct name to apply.” That comment in 1957 “launched

me into the taxonomy and the nomenclature of the fungi” Grégoire

became an exceedingly meticulous collector, who knew the importance of

thoroughly screening a potential type locality and delivering a reliable type

specimen: “Later wanting to provide sufficient type material for herbarium

deposit [of Arachnophora fagicola] in view of publication, I had to examine

xiv ... MYCOTAXON 136(4)

4. Verrucobotrys, 5. Dichobotrys; 6, Chromelosporium; 7. Pulchromyces; 8. Phymatotrichopsis; 9.

Ostracoderma; 10. Glischroderma.

Fic. 2. The many faces of Botrytis sensu lato.

eight thousand beech cupules from the same locality to finally have eighteen

minute colonies of that new fungus to serve as a type.”

Of the 380 Botrytis names in the literature, many were already synonymized

while others were described through short Latin descriptions frequently lacking

illustrations. Grégoire turned to nature: “exploring woods, swamps, ponds,

fields, vineyards, orchards and vegetable plots, flower gardens, agricultural and

forest soils, even caves ... collecting specimens and [isolating] Botrytis strains

in pure culture.... [Most] cultures were conidial and sclerotial. I was disturbed

finding in some of them small globose conidia—like Penicillium conidia—even

in single Botrytis conidium cultures. I could not obtain their germination and

concluded that they were spermatia....”

Realizing that these Botrytis forms were asexual and ‘imperfect; Hennebert

returned to the field to collect apothecia on sclerotia together with Botrytis

conidiophores (representing Botryotinia globosa, B. ficariarum, and B. calthae)

under the host plants Allium ursinum, Ficaria verna, and Caltha palustris.

The apothecia, produced by sexual ascospores, represented the sexual (‘perfect’)

states of the asexual Botrytis stage.”

The spontaneous development of apothecia in a Botrytis porri pure culture

(isolated from Allium porrum) raised the problem of nomenclature: “To each

state of the fungi, imperfect state and perfect state, a distinct Latin binomial

OCTOBER-DECEMBER 2021... XV

name was given. This is heritage of the past when mycologists considered each

reproductive form a distinct species.... Several names for one fungus species

appeared contrary to Linnés principle and that of the International Code

of Botanical Nomenclature, one taxon—one name. In such cases the Code

prescribes the predominance of the name of the sexual form over the other

names [but] tolerates ... the use of the other names. That peculiarity of the

nomenclature of the pleomorphic fungi struck me....

“Soon I had hundreds of Botrytis isolates in pure culture beside isolates of

many other fungi, an embryo of a fungus culture collection or MYCOTHEQUE....

‘But why bring nature in small tubes?’ questioned a relative of mine, long

ago. Why a fungus culture collection? Why a mycotheque?’ [I replied] ‘to

help elaborate a more natural taxonomy of the fungi, through demonstration

and comparison in pure culture of their morphology and pleomorphy, and

correlatively to resolve nomenclatural problems that pleomorphy may cause.

TAXONOMY and NOMENCLATURE were, to me, the prime reasons for building

a mycothéque.”

In 1995 Grégoire summarized, “Taxonomy and nomenclature are at the base

of all activities of man who observes, thinks, and communicates. Observation

leads to thought, thought to concepts. Taxonomy is the result of observation

and thought. Nomenclature results from thought and concepts. Taxonomy and

nomenclature [are] at the base of all common knowledge.... Observation of the

diversity of things induces comparison and distinction and leads to elaboration

of concepts and to formation of words. If a concept is a category, the word

is a noun; if it is an attribute, the word is an epithet. Categories are genera,

epithets refer to species. That is taxonomy. Nouns are generic names, nouns

plus epithets are species names. That is nomenclature. Common knowledge is

made of taxonomy and nomenclature’

During 1962-65 Grégoire was appointed UCL Plant Pathology Teaching

Associate and charged with developing the Fungus Culture Collection

N how do Ow wot

Vere pege A S labelled?

in Heverlee-Leuven. RIGHT: Taxonomy vs nomenclature in the ‘older’ days.

XVI ... MYCOTAXON 136(4)

—s> Genealogy of the MUCL culture collection Coie

1934 1994-1995-1997 2006

weeeeeeeeeeen Y eee een }=Co ooo POCO Foo ooo Pos

1894 1968 1999

* Purification and selection of brewery yeasts

* 1894; public inauguration of the Biourge Culture Collection

* 1898-1901: taxonomic study of Penicillium by F. Dierckx

nn , * 1901: vitamin “Bios” discovery by E. Wieldiers

Philibert Biourge O- 1894 * 1923: Penicillium monograph with 126 species

* 1927: monograph of the genus Aspergillus

* 1929: industrial production of citric acid from Aspergillus niger by A. Cappuyns

* 1932: studies on the Aspergillus group niger by R. Mosseray

7 O * 1939: discovery of Griseofulvin; Mycothéque Philibert Biourge

Paul Simonart 1934 * Setup of quality control for dairy products with R. Lambert

V * 1968: Mycothéque de I’Université catholique de Louvain (MUCL)

* 1969: integration of an African soil Hyphomycete collection by J. Meyer

¢ 1972: affiliation of MUCL to WFCC (World Federation for Culture Collections)

* 1977: introduction and definition of the terms anamorph, teleomorph and

holomorph with L. K. Weresub

Grégoire L. Hennebert O-— 1968 > 1983: launch of BCCM (Belgian Co-ordinated Collections of Micro-organisms);

ECCO membership (European Culture Collections’ Organisation)

* 1989: first edition of the MUCL catalogue of fungi and yeasts

* 1992: recognition of BCCM as an IDA (International Depository Authority) under

the Budapest Treaty

vV * 1994: celebration of the MUCL Centenary

1994 —O Jacques Decallonne

1995 -—O Anne-Marie Corbisier

1997 —O _ Tomas Avella-Shaw

* 2001: participation in European programs (e.g. COWIDI and ESA);

development of GINCO (Glomeromycota in vitro collection) by S. Declerck

Henri Maraite O-— 1999 - 2002: strengthening of cryopreservation infrastructures

* 2005: ISO 9001 certification of the collection’s core activities; first international

training on in vitro culture of AMF by S. Declerck

vV * Strengthening of industry partnership; anchorage in European programs

{e.g. EMBARC, BRIO, VALORAM, ERAFRICA)

* 2012: development of cryopreservation methods for recalcitrant strains and

microbial consortia

Stephan Declerck O- 2006 ° 2014: setup of collection, research, service and communication strategies

* 2015: go-live of the LIMS (Laboratory Information Management System)

* 2016-2017: re-localisation of the cryopreserved backup collection

* 2017: first BCCM training course on preservation of micro-organisms

Vv * 2017-2021: orientation towards microbiome research

pdf (https://baccm.belspo.be)

| M4 Earth and Life Institute

Agro-food|& Environmental Fungal Collection Mycology - Mycothequeide I/UGE U L

Groix(dulSud) 2!box/L7105:06 Université

4348 louvain:la:Neuve = BELGIUM catholique

de Louvain

BEL'GIAN|GO-ORDINATEDIGOLLEGTIONS OF MIERO-ORGANISMS

BCCM/GREAT/AT SMALE THINGS

becm:belspo:be

beem-mucl@uclouvain:be

Tel:)+32:10/47 37/42

MUCL_ Timeline.

Fic. 4. The 1894-2021 MUCL time-line

at the Faculty of Agriculture, Heverlee-Leuven (Fic. 3); this preceded his

eventual development of Mycotheque de l'Université Catholique de Louvain

[MUCL]; during his 26-year tenure as supervisor of his beloved mycotheque,

he guided the expansion of the 300-strain culture collection from its modest

OcCTOBER-DECEMBER 2021 ... XVII

beginnings as the Mycotheéque Biourge to 35,000 fungal strains and (by 2010)

the curation of 51,000 dried specimens (Fic. 4).

Courtes) of Gertrude Hennebert

Fic. 5. Young Grégoire at the lab bench at the Catholic University of Louvain (date unknown).

Grégoire matriculated at the University of Louvain (UCL) in 1947, receiving

his bachelor’s in philosophy as Engineer in Tropical Agriculture and

becoming a Superior Scientific Education Graduate Associate in African

Sciences. His four years researching Plant Pathology at UCL were funded by

the Fonds National de la Recherche Scientifique.

After successfully defending his PhD thesis on Botrytis and the

Sclerotiniaceae in 1960, Grégoire was awarded a 2-year postdoctoral

Fellowship with Dr. Stan Hughes of the Plant Research Institute in Ottawa,

Canada, a time that he recalled with great fondness, and which re-introduced

him to the joys of field work (Fic. 6). “The first step of any mycological study

is the observation of the fungi in the field. To teach this basic requirement,

Dr Stanley J. Hughes, the first day after my arrival in Canada for a two-year

postdoctoral stay (1960-62), took me to the forest, in Gatineau Park, to collect

microfungi, field shoes on feet, knife and basket in hands.”

“Professor Richard P. Korf, who I visited at Cornell University in 1961 for

the first time of many, was doing the same with his students: ‘no mycology in

Xvitl ... MYCOTAXON 136(4)

Fic. 6. THE IMPORTANCE OF FIELDWORK. LEFT: Prof. Stanley Hughes collecting in Gatineau

Park, near Ottawa in 1960. RIGHT: Prof. Richard Korf in 1961 sorting fungal specimens during

a field trip near Ithaca, New York.

class or laboratory before a discovery of the fungi in the field’ The bonnet of

the car could serve as a bench to sort the collections. But take care: some fresh

polypores, like Ganoderma applanatum, exude drops of enzymes so active that

within one hour's time you may have to ... repaint the bonnet.”

Grégoire became Professor in 1971 and registered MUCL with the WFCC

and IAPT in 1972. During 1964-75, he was a regular scientific collaborator of

the Centraalbureau voor Schimmelcultures, visiting Baarn every two weeks.

In 1975, after being “obliged to move laboratory, herbarium, the MUCL

culture collection, and furniture 30 km south of Brussels onto the bare new

campus named “Louvain-la-Neuve,’ he was elected president of the Faculty of

Microbiology and Bio-industries, a position he held for five years.

Throughout these decades Grégoire served as a visiting lecturer/professor

throughout North America, Africa, and Europe at Cornell University (Ithaca,

New York), the University Lovanium in Kinshasa (Congo; 1967), Québec’s

Laval University (1970), Prague's Czechoslovakian Academy of Sciences (1970),

South Africas Potchefstroom University (1981), Bujumburas University of

Burundi (1982-90), Slovenia’s Boris Kidric Institute of Chemistry (Ljubljana),

and Morocco’ Institut Agronomique et Vétérinaire d’Agadir (1986). During

this period was also invited by Prof. A.M. Cotes-Pradohe to evaluate the

CORPOICA Germplasm Bank in Bogota, Colombia.

Grégoires African tropical research revealed some problems associated

with following a strictly classical approach when faced with largely unexplored

fungal diversity. These issues formed the basis for a spirited discussion among

the 150 registrants of the 1994 MUCL Centenary Symposium (Fic. 7). “A

priority and urgent duty is to describe and make the inventory of this mycoflora,

OCTOBER-DECEMBER 2021 ... XIX

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Mid-l&r: courtesy of Gertrude Hennebert

Fic. 7. THE 1894-1994 MUCL CENTENARY SYMPOSIUM—A TIME FOR SCIENCE & REUNION.

Top: Official conference photo, 29 June 1994. mip-LErT: In 1974 C.V. Subramanian (hiding two

steps above Grégoire in the top photo and who first visited MUCL in 1965) receives a handshake

from Grégoire in India congratulating him on his 50" birthday. mip-RIGHT: Grégoire & Stan

Hughes at the microscope during Stan’s 1973 visit to MUCL. BoTTOM LEFT: Stan Hughes and

Dick Korf at the 1994 symposium. BOTTOM RIGHT: Lidwina Hennebert (left, next to Grégoire’s

empty chair) and Dick & Kumi Korf (right) at the symposium dinner at the Abbey Hétel of

Villers-la-Ville.

XxX ... MYCOTAXON 136(4)

using microscope and culture in situ, for which well-trained young mycologists

are needed. And on that basis, elaboration of monographs should provide the

necessary tools to further investigations.” The symposium attendees agreed

that tropical fungi differ from those of the temperate regions and that morpho-

anatomical taxonomy had to be complemented by molecular taxonomy.

“Good phylogeny based on molecular data may demonstrate relationships

that might lead one to amend the system of the fungi at the highest ranks.”

They agreed that a new training profile was needed for tropical and temperate

regions covering ... “different disciplines, including computer science, cladistics,

molecular biology, [and] ecophysiology without neglecting ... traditional

but unavoidable and basic methodologies of fungal taxonomy—collecting,

microscopying, drawing, isolating fungi into pure culture, studying type

material, interpreting past literature, writing descriptions, constructing keys,

writing monographs, building expert systems of identification and managing

herbaria and culture collections. These are the first acts for a mycologist in

tropical and unexplored areas of the world.”

FounDING Mycotaxon: In 1960 at the beginning of his post-doc with Dr.

Hughes in Ottawa, Grégoire met Dr. Richard P. Korf for the first time. “Dick

invited Lidwina and me to his home along the Lake in Ithaca in the summer

1961. At his door, when we arrived, we found the message “Welcome, come in,

the house is yours, we come very soon, Dick’ Surprising!” During September

1969 (at the Kananaskis Conference on pleomorphy) “we discussed [day and

night] the nomenclature of the pleomorphic fungi [and] about my distinction

between botanical and anatomical nomenclature. I was greatly honored when,

as Fulbright Scholar in 1972-73, he chose to stay with us at the University of

Louvain.” (Hennebert 2018)

Dick had visited MUCL on 13 June 1966. During his 6-month-long stay

in 1972-73, in addition to hosting a conference on ‘A new classification of the

operculate discomycetes, he discussed with Grégoire the need for a timely

mycological publication geared to taxonomy and nomenclature. “Even after

searching for guarantee of the novelty, it is important to publish rapidly.

As very few journals were editing manuscripts with a delay shorter than 6

months, [Korf] and I decided to edit a new journal offering only a short delay

together with quality, designed to promote the taxonomy of fungi and lichens”

(Hennebert & Korf 1974).

Their proposed endeavor turned out to be a life-time commitment.

Since sending out its first volume in July 1974, MycotTaxon has published

OCTOBER-DECEMBER 2021 ... XXI

Fic. 8. HENNEBERT & KORF

at Louvain-la- Neuve making

plans during Korf’s 1972-73

Fulbright studies to launch

a new mycological journal

devoted to the taxonomy

and nomenclature of fungi

| and lichens.

136 volumes comprising approximately 82,000 pages in its journal. In 2004,

Dick, Grégoire, and the Editorial Board agreed that we remove the overly

lengthy mycobiota (annotated regional species lists) from the journal to be

summarized/abstracted in the journal and posting ~3500 much larger pages

for free download from the MycoTaxon website. The founders and Board also

approved (in 2004) moving to editorially generated copy, (in 2011) offering both

online and hard copy subscriptions, and (in 2014) moving to annual volumes

comprising four quarterly issues. While Grégoire and I mourned somewhat the

move to online publishing (I chose black for the cover announcing the change),

Dick welcomed it as a positive “green” solution.

Dick served as Managing Editor & English Language Editor (1974-91),

Assistant to the Editor-in-Chief (1998-2001), Business Manager (1991-

2006), and Treasurer until shortly before his death in 2016. Grégoire served

17 years as Book Review Editor (1974-91) and 32 years as French Language

Editor (1974-2006). His many years of service to the journal continued until

the end: he published in Mycotaxon his last three papers on Glischroderma

XXII ... MYCOTAXON 136(4)

(Hennebert 2018), the possible congeneric relationship between Pachyphlodes

and Chromelosporium (Hennebert & Decock 2020), and a reevaluation of

Chromelosporium and description of two new genera Chromelosporiopsis &

Geohypha (Hennebert 2020).

ENTER ANAMORPH, TELEOMORPH, AND HOLOMORPH: In his last lecture,

Grégoire noted that the diverse conditions in culture permit ‘the fungus to

reveal itself in more facets than expected,” thereby helping demonstrate the

variability and mutability of a fungus.... Only through single spore culture,

[could it] be demonstrated that spermatia belong to Botrytis cinerea.”

Linnaeus (1753), who based his taxonomy on sexual reproductive structures,

classified plant taxonomists according to their basic principles of classification,

considering himself a ‘sexualist;’ with fungal sexuality and pleomorphism

unknown in the 18" century, any distinct reproductive form was considered a

distinct species and named (leading inevitably to a proliferation of names for a

single species). Grégoire’s early study of conidiogenesis in culture had reduced

the taxonomy of Botrytis to some 36 species but also underscored the confusion

surrounding life stage nomenclature.

Tulasne, who in 1851 called pleomorphy the coexistence of several forms of

reproduction in fungi, demonstrated the organic connection between ascosporic

fungi and conidial fungi as the ‘perfect [or sexual] state’ ... and the ‘imperfect

state or states’ of the same species, as several conidial forms of reproduction

might exist in the same ‘perfect’ species. Eventually Fuckel classified fungi as

belonging to the Fungi Perfecti and the Fungi Imperfecti. Through his 1867

LOIS DE LA NOMENCLATURE BOTANIQUE Art. 15, De Candolle “reinforced the

principle ‘one taxon—one name, while Saccardo (1904) recommended a dual

nomenclature for pleomorphic fungi in Fungi Perfecti and Fungi Imperfecti,

thereby supporting usage of ‘imperfect’ names beside the ‘perfect’ name.

The 1910 Brussels Botanical Congress integrated Tulasne and Saccardos

positions, recognizing for ascomycetes and basidiomycetes the predominance

of the name of the perfect state over the names of their imperfect states, thereby

tolerating the use of the existing names of imperfect states to be classified in

‘form-genera’ and assigning them ‘only a temporary value:

In 1966 the Edinburgh Code established Art. 59, intended to separate

the imperfect and perfect nomenclatures while preserving the stability of the

names of perfect species. Korf (1982) noted that in practice, however, Art. 59

was “undoubtedly one of the thorniest articles of the Code for mycologists to

apply,’ and Hawksworth (1984) deemed it “extraordinarily complicated, so that

OCTOBER-DECEMBER 2021 ... XXIII

in some instances, up to three separate conclusions could be drawn regarding

names to be used.”

At the 1969 Kananaskis Conference on Taxonomy of the Fungi Imperfecti,

Grégoire presented a paper on the fundamentals of pleomorphic and

pleoanamorphic fungi, in which he (supported by Korf) formulated the

concepts of ‘phase’ vs ‘state; ‘form, ‘whole fungus’ vs ‘perfect’ & ‘imperfect states,

‘botanical taxon’ vs ‘form-taxon, and ‘botanical’ vs ‘anatomical’ nomenclature in

order to define adequately the divergent options applied by fungal taxonomists

in naming pleomorphic fungi (Hennebert 1971).

In 1969, Grégoire became a member of the International Association

of Plant Taxonomy’s Special Committee for the Nomenclature of Fungi and

Lichens (IAPT-N), which he served until 1986, and was one of 12 members

in Subcommittee A at the First International Mycological Congress in Exeter,

September 1971.

He recalled that that in 1977, “just playing with words in a conversation

with Dr Luella Weresub [the formidable chair of Subcommittee A whom he

knew from his 1960-62 studies in Ottawa], I proposed some less ‘imperfect’

terms than the anthropomorphic ‘perfect’ and ‘imperfect; hopefully terms

without value judgements ... to designate the anatomical parts of a fungus

deserving a name.... We published the terms ‘anamorph, ‘teleomorph, and

‘holomorph to differentiate any fungal ‘morph’ deserving a name (Hennebert

& Weresub 1977) .... The term ‘anamorph’ was proposed for any asexual state

of the [asexual phase], the term ‘teleomorph’ for the unique sexual state of the

Fic. 9. LUELLA WERESUB was the first woman faculty

member to be hired in the Department of Botany of

the University of Manitoba in 1952, moving to the

mycology unit of the Department of Agriculture in

Ottawa in 1957, where she worked until her death

in 1979. Dr. Weresub, an expert in corticioid fungi

and sclerotium-producing basidiomycetes, became

a world authority in the botanical nomenclature

of fungi and particularly influential member of

the IAPT Nomenclature Committee for fungi and

lichens as Chair of the Subcommittee on Article 59.

https://www.facebook.com/umanitobasci/

XXIV ... MYCOTAXON 136(4)

fungus, and the term ‘holomorph’ for the ‘whole fungus’ (which encompassed

all unknown ‘morphs’).”

This proposed modification of Art. 59 was recommended in 1977 by

the Second International Mycological Congress in Tampa and adopted by

the Sydney Code (Stafleu 1981). While simplifying the previous rule, the

language reinforced segregation of ascomycete and basidiomycete anamorphic

nomenclature from teleomorphic (holomorphic) nomenclature based on their

‘anatomical’ typifications, clearly declaring “anamorphic names unpriorable

in holomorphic nomenclature in case the anamorphic fungi were ever found

to be sexual. It was the most conservative option, far from the true botanical

nomenclature requiring the suppression of Art. 59, [which] I considered the

final solution.” The new Art. 59 did offer two advantages: it solved “a serious

nomenclatural knot by a clear practical line” and stimulated reconsideration of

the basic principles of fungal nomenclature.

“For plant pathologists, anamorphic names of plant pathogens often serve

to designate plant diseases. Similarly, in medical mycology, names of mycoses

commonly derive from the anamorphic name of the causal agent. Both practices

are regrettable since they link disease names to a not infrequently changing

reference basis. ... The true reason [for] maintaining Art. 59 was the fear of the

major disturbance that its suppression and a return to unrestricted typification

and absolute priority of names would cause ....

“Obviously, good sense must prevail. The fundamental aim of naming living

beings is indeed to make possible their recognition, inventory, and classification,

by giving each of them a single name and nota multiplicity of names (Hennebert

1991). I therefore prefer giving one name only to pleoanamorphic fungi. So

I proceeded in naming and describing Botrytis, Chalaropsis, Wardomyces,

Mammaria, Humicola species, using eventually a cross-reference generic

denotation for what I considered the ‘secondary’ conidiogenesis.”

In his memoriam to Grégoire in IMA Fungus, David Hawksworth observed

that it was “gratifying to see how [Grégoire] came to openly accept the demise

of the use of the ‘-morph’ terms when the separate naming of morphs of

pleomorphic fungi was abandoned in 2011. This is evidenced in his last

publication ... where he revisited and revised some of his previous taxonomic

and nomenclatural conclusions (Hawksworth 2021).

GREGOIRE LAURENT HENNEBERT was born on 20 June 1929 to Dr. Raymond

Hennebert and Marie (Wallez) in Mons, Belgium (Fic. 10). In 1960 he

married Lidwina Baert, with whom he shared a long and happy marriage

=9053

http://www.paulcuvelier.Org/?p

Courtesy of Gertrude Hennebert

OcCTOBER-DECEMBER 2021... XXV

| t

“oy

Fic. 10. THE Dr. HENNEBERT FAMILY IN Mons, BELGIUM, 1944. LEFT TO RIGHT: Grégoire (age 15),

Alexis, Dr. Raymond Hennebert (father), Paul, Julie, Marie Wallez Hennebert (mother), Colette.

Fic. 11. Grégoire remained active during his

‘emeritus’ years. Daughter Gertrude writes:

“In his spare time, my father was a passionate

beekeeper. He loved to share his passion with

his grandchildren. In these two pictures you

can see him in action.”

XXVI ... MYCOTAXON 136(4)

until her death in 2009, and was the proud father of four—Gertrude,

Mechtilde, and Imelda Hennebert and Khan Nuhr—and grandfather to

twelve, many of whom he taught about the joys of beekeeping (Fic. 12).

Grégoire exercised regularly and was in unusually good health, expecting

to reach 100 after his successful heart surgery in 2021. Unfortunately,

after falling in his garden and breaking a vertebra in his back, he was

hospitalized and rapidly declined, slipping away five weeks later on

13 October 2021. His daughter Gertrude wrote that Grégoire would “never

have thought that he would die as the consequence of a fall.”

Honored by the genus Hennebertia and seven hennebertii species epithets

(in Acremonium, Chaetopsis, Cylindrotrichum, Gliocladiopis, Niesslia,

Penicillium, and Raffaelea), Grégoire himself named 18 genera and 114

species. He guided 55 research studies in applied mycology (26 at the

University of Burundi) and published alone or with colleagues 156 scientific

papers (not including 13 culture papers). He also edited the 100 YEARS OF

THE FUNGUS COLLECTION MUCL 1964-1994 symposium proceedings, two

books on Les Mérules des Maisons, and the Directory of African Mycology

and the Aspects of African Mycology from the first Regional Conference on

Mycology in Africa.

Grégoire remained mycologically active after his retirement in 1995,

although he did not have particularly fond memories of his ‘emeritus’ status. In

2017, he explained that the long delay in preparing his paper on a Glischroderma

collected by Dick in 1994 arose from problems caused by a misunderstanding

over his taking the books reviewed for MycoTAxon (contractually his) and the

expectation that he administer the lab without pay for a year, after which he was

denied lab access. Not until he discovered Dick's potential type in Geneva, did

he remember his unfinished paper. Although fairly certain that the specimen

represented a second species, he described it as Glischroderma aff. cinctum.

“Fresh collections and DNA analyses might solve the question better than I can

do. I think I shall just raise the question. I shall not be ‘waiting for that elusive

muse, perfection, although it is a temptation...”

On discovering a nonsensical but rather major error in the final proof of his

2020 paper with Cony Decock that none of us had noticed in previous versions,

Grégoire became understandably distraught. My prompt repair elicited this

grateful note, “Dear Lorelei, Many thanks for the exceptional correction, as a

New Year gift. It is now perfect. Iam happy and makes me better in insomnia

and allows me to go to sleep again.”

OCTOBER-DECEMBER 2021 ... XXVII

Courtesy of Gertrude Hennebert

Courtesy of David Hawksworth

Fic. 12. Post ‘retirement life. LEFT: Peter Austwick, Grégoire, and Stan Hughes relaxing during

the 1996 Centennial Meeting of the British Mycological Society in Sheffield, England. r1GHT:

Grégoire and Walter Gams discuss Article 59 and the launching of MycoBank at the Centenary

of the Centraalbureau voor Schimmelcultures in Amsterdam in 2004.

His 2018 Glischroderma paper led him to reexamine his Chromelosporium

archives and the monograph he began with Stan Hughes in 1960-62 while in

Ottawa but had never completed despite encouragement from both Stan and

Dick. Feeling he owed it both departed friends that he complete the work,

Grégoire also felt it necessary to sequence as many specimens as possible to sort

genera and nomenclature correctly. “With the precious help of Rosanne Healy

and MUCL, I progress in the revision of the genus, including nine and possibly

more species. But it progresses slowly with so many other things needing to be

done.... 1am sorry to inform you that on 21 December [2017] I got a heavy car

accident stopping against a tree. I have nothing broken but the chest smashed

in by the belt, making the holidays painful. I wish you a happy and healthful

New Year.”

Grégoire prepared his final paper under pandemic conditions. Replying

to my concern about the lock-downs and huge numbers of Covid-19 cases

sweeping his country, he noted, “Yes Belgium is in bad health situation by lack

of foresight. From begin of March, the virology laboratories were all lacking

chemicals for testing, while the virus was already present. And the stay-home

rule was decided only on March 15 [2020]. I stay home and have visit of my

daughter only when needed.... You should be safe in your Nature environment.

I have only a piece of ground of 30 x 50 m with my house, a piscine, and some

green (now absolutely brown by the drought) but sufficient to be far enough

from neighbours.”

I never had the pleasure of meeting Grégoire in person but came to

greatly enjoy our correspondence. Meticulous as he was, Grégoire worked his

reviewers (Kathy Hodges, Roseanne Healy, Keith Seifert, David Hawksworth)

XXVIII ... MYCOTAXON 136(4)

Courtesy of Gertrude Hennebert

Fic. 13. Grégoire Hennebert turns 80 (2009).

and editors exceedingly hard, yet received all critiques graciously, expressing

his gratitude frequently. “Dear Lorelei, I just received my last paper reviewed

by Keith Seifert. He proposed many amendments, a lot of work for me”

Extremely worried that his ‘last’ paper would not get published, he greatly

appreciated that David and Keith came to his aid. Shaun and I, who spent

well over 100 hours emailing, revising, and adjusting, were rewarded by his

immense relief when it was released in MycoTaxon 135(3). On 9 October

2020 he wrote: “I am very pleased with the paper, and I thank both Shaun and

you so much for the improvement of the manuscript, the time you putted in

it, and the priority you gave it by agreeing to publish it in the year 2020. With

loves, Grégoire” This was followed on 30 October 2020 by, “Dear Lorelei, dear

Shaun, I just received MycoTaxon 135(3) via Lulu and found my ‘last paper’

in full on 54 pages and a kind comment by yourself in the introduction of the

volume. I really want to express all my thanks to Shaun and to you for the

edition of my paper in MycoTAxon 2020. Now, although there are some other

unachieved projects (which are for others), I conclude by this paper my work

in Mycology with real satisfaction. With many thanks, Grégoire.”

Grégoire was a warm and kind correspondent, frequently sharing advice

gained from the successful cancer treatment and preventative measures

OCTOBER-DECEMBER 2021 ... XXIX

(immunotherapy, shiitake, Grifola frondosa, Cordyceps militaris) used after

Lidwina’s bout with breast cancer. A believer in healing touch therapy, he also

used willow herb (Epilobium parviflorum) for his own prostate cancer and

hawthorn berry and heather teas to help him heal after his 2010 and 2012

heart attacks. Beyond doubt, his positive attitude kept him healthy for so long,

“Absolutely YES, Life is good and beautiful. I enjoy it fully”

Gregoire was declared Professor Emeritus after delivering his ‘last lecture’ on

5 December 1995, yet he published his last papers in the journal he co-founded

almost a quarter century later. We are grateful that he and Dick established

MycotTaxon 48 years ago and will continue to miss our two mycological giants,

whose vision was always for the future.

—Lorelei Norvell

Fic. 14. GREGOIRE HENNEBERT in his garden,

smiling for his 90" birthday photo.

Courtesy of Gertrude Hennebert

ACKNOWLEDGMENTS

We greatly appreciate the many photos and information generously provided

by Grégoire’s daughter, Gertrude. Unless otherwise noted, quoted text was taken

from Grégoire’s final 1995 lecture, private autobiographical notes, and MycoTaxoNn

editorial correspondence. All unattributed photographs were generated from

computer screen shots of illustrations in the PDF version of THE 100 YEARS OF

THE FUNGUS COLLECTION MUCL 1894-1994 (Hennebert 2010), covering both

the proceedings of the symposium attended by 150 registrants on 29 June 1994 at

the Catholic University at Louvain-la-Neuve and Grégoire’s complete final lecture,

delivered on 5 December 1995 prior to his becoming Professor Emeritus.

Xxx ... MYCOTAXON 136(4)

SELECTED BIBLIOGRAPHY

Hawksworth DL. 2021. Grégoire Laurent Hennebert (1929-2021), p. 32 in MycoNews. IMA

Fungus 12: 1-45. https://doi.org/10.1186/s43008-021-00085-9

Hennebert GL. 1960. Recherches morphologiques sur le genre Botrytis Persoon. These Univ. Cath.

Louvain, Inst. Agron. Heverlee. 188 p.

Hennebert GL. 1971. Pleomorphism in fungi imperfecti, 202-223 in Taxonomy of fungi imperfecti

(WB Kendrick, ed). Univ. of Toronto Press. 309 p..

Hennebert GL. 1991. Art. 59 and the problem with pleoanamorphic fungi. Fourth International

Mycological Congress, Regensburg, 28 August-3 September 1990. Mycotaxon 40: 479-496.

Hennebert GL. 2010. Taxonomy and nomenclature of the fungi: reasons for a Mycotheque [Last

lecture: 5 December 1995]. 257-302, in GL Hennebert (ed.), The hundred years of the fungus

collection 184-1994. Mycotaxon Ltd. (see directly below).

Hennebert GL (ed.). 2010. The hundred years of the Fungus Collection MUCL 1894-1994—fungal

taxonomy and tropical mycology: Quo vadis? Taxonomy and nomenclature of the Fungi.

Mycotaxon, Ithaca. 318 p. [available on www.mycotaxon.com]

Hennebert GL. 2018 (“2017”). Glischroderma Fuckel. Mycotaxon 132: 745-757.

https://doi.org/10.5248/132.745

Hennebert GL. 2020. Chromelosporium re-evaluated, with Chromelosporiopsis gen. nov. and

Geohypha stat. nov. Mycotaxon 135: 665-718. https://doi.org/10.5248/135.665

Hennebert GL, Decock C. 2020. A comparison of anamorphs of some Pachyphlodes species and

the type of Chromelosporium: are they congeneric? Mycotaxon 135:167-182.

https://doi.org/10.5248/135.167

Hennebert GL, Korf RP. 1974. Mycotaxon, a new international journal on taxonomy and

nomenclature of fungi and lichens. Mycotaxon 1(1): 1-12.

Hennebert GL, Weresub LK. 1977. Terms for states and forms of fungi, their names and types

Mycotaxon 6(1): 207-21.

From Ammirati & Glawe (2008)

OCTOBER-DECEMBER 2021 ... XXXI

IN REMEMBRANCE— Jack RoGers (1937-2021)

Fic. 1. Professor Jack D. Rogers at 80 standing

beside the Department of Pathology display case

at Washington State University in Pullman.

MycoTaxon mourns the loss of a second mycological giant and friend,

the longtime Washington State University professor and internationally

renowned mycologist famed for his “booming voice, distinctive West

Virginia accent, disarming and endearing sense of humor, enthusiasm for the

outdoors, and genuine care for students and colleagues.” Jack DaviD ROGERS

died at home in Pullman, Washington on June 14 at the age of 83. Deep

condolences go out to Belle, his wife of 58 years, twin daughters Rebecca Ann

(Hines) and Barbara Lee (Cooper), and family. This brusque but lovable man

will also be greatly missed by his international network of colleagues, friends,

and former students.

Professor “Xylariaceae’ was born 3 September 1937 in Point Pleasant, West

Virginia, to schoolteachers Jack Rogers (Youkobis Uzskuriatis) and Thelma

(Coon) Rogers. Young Jack graduated from Point Pleasant High School in 1955,

earned his BS from Davis & Elkins College (WV), marrying fellow student Belle

Clay Spencer in June 1958, his junior year. After obtaining his masters in Forest

Management from Duke, he earned his PhD in 1963 under the supervision

xxx ... MYCOTAXON 136(4)

of John Berbee at the University of Wisconsin-Madison for his research on

Hypoxylon pruinatum (= Entoleuca mammata), a major forest pathogen of

quaking aspen in the Great Lakes States.

Directly after receiving his doctorate, Jack joined the College of

Agriculture faculty of Washington State University, where he was to

remain for the rest of his 50-year educational career. Appointed to

Assistant Professor in the Departments of Plant Pathology and Forestry

& Range Management, he quickly advanced to the rank of Full Professor.

Throughout his university tenure, he taught classes in mycology and forest

pathology, frequently coordinating departmental seminar series. As part

of his advanced mycology series, he developed the Ascomycetes and Fungi

Imperfecti course, which he taught for 36 years, and which persuaded many

students to pursue mycological careers. He helped rework the Pathology

Department curriculum to offer Advanced Fungal Biology, also introducing

a popular general studies course—Molds, Mildews, and Mushrooms: The

Fifth Kingdom. During 1985-99, Jack served as Chair of the Department

of Plant Pathology while maintaining a full teaching schedule and active

research program and authoring/co-authoring 70 scientific papers. Eleven

master’s and 17 doctoral students earned their degrees under his guidance.

Outside of family and fishing, Jack’s passion was teaching, research, and

the university. Washington State reciprocated by recognizing his educational

and scientific excellence through many awards: its 1967 R.M. Wade Award for

Instruction; 1986 Sahlin Faculty Excellence Award for Research, Scholarship,

and Arts; 2005 Library Excellence Award for Service to WSU Libraries; and

2006 WSU Eminent Faculty Award. Jack was promoted to Regents Professor

in 2007 and retired from the WSU faculty in 2013 after 50 years of devoted

and exemplary service.

Jack definitely had presence, even when proudly holding a steelhead

(Fic. 2). Given his reputation as a humorous raconteur, he was a surprisingly

formal instructor. Dr. Brenda Callan, student and fellow Xylaria expert,

writes, “he required all of his students, including graduate students, to address

him formally, as ‘Dr’ Rogers. Sometime after I completed my Ph.D., he gave

me permission to address him by his first name. It was a difficult transition,

rather like starting to call one’s own father by his first name.” Brenda (as have

all of us who knew Jack) remarked on his deep voice: “Who could think of

Jack and not hear in their mind his voice, that booming drawl? My desk

was in the herbarium, and his classroom around the corner and several

doors down. I never had to peek into the classroom to see if it was time

OcCTOBER-DECEMBER 2021 ... XXXIII

to go in to help with the labs. He always seemed surprised when I showed

up right after hed finished lecturing!” Dr. Callan also commented on Jack's

unorthodox morphological descriptions, “Jack’s prose sometimes painted

a vividly colourful, oddball image of the fungus he was describing. For

example, in Rogers & al. (1987), he describes the stromata of Kretzschmaria

cf. mauritanica: “The surface of our material resembles a deeply cracked

chocolate cookie that has been dusted with sugar’ I think that was written

close to lunch time.’ (Callan, pers. comm.)

WSU Plant Pathology Chair Dr. Timothy Murray knew Jack for 43 years,

first as a graduate student and then fellow faculty member. He told the

Washington State Insider covering Jack’s death that “I never worked harder

in a class than when I took his class.... He really was the complete package:

He was a great scholar. He won awards for his research. He was very prolific

in terms of publication. He was an engaging and committed teacher. And

he worked on a number of higher-level university committees. He loved all

aspects of the job, and he did them well.... He was also ‘a real character. He

loved to tell a good joke. And he was very vocal in terms of explaining his

position. If he had an opinion and it differed from yours, you would know

what his opinion was.” (Janovich 2022)

Even after his 2013 retirement, Rogers continued coming to his office. Dr.

Lori Carris, his former master’s student, Tilletia expert, colleague, and now

WSU Professor Emerita, noted, “It was kind of a challenge to see if I could

get there before Jack.... The only time he took off consistently was Wednesday

afternoon. That's when he went fishing.... He loved being out in the woods.

He loved anything to do with fungi” (Janovich 2022)

Jack did not restrict himself to his own university but was very active

in the Mycological Society of America (MSA), which he served as Western

Councilor (1970-72), Vice President (1975-76), and President (1977-78),

eventually receiving its two most prestigious awards, the Weston Award for

Teaching Excellence in 1992 and Distinguished Mycologist Award in 2004.

It is no surprise that fellow ascomycetologist Dick Korf enticed Jack onto

Mycotaxon’s first Editorial Advisory Board, which he served well for eight

years (1990-98).

An international authority on xylariaceous fungi, Rogers authored and co-

authored more than 230 scientific papers and two mycological books. We

have tallied one family (Graphostromataceae), 22 genera, c. 240 new species,

35 new varieties, and c. 150 new combinations named or co-authored by Jack

XXXIV ... MYCOTAXON 136(4)

during his career. In 2017 Wendt, Kuhnert & Stadler honored him with the

genus Jackrogersella.

Rogers traveled the world for research, collecting specimens and helping

curate WSU’s Charles Gardner Shaw Mycological Herbarium [WSP]. The

National Science Foundation funded his research for more than 30 years,

which, Dr. Carris noted, “really laid the foundation for a lot of the work on

Xylariaceae that is going on today.” Jack was an especially engaging collecting

companion. Aware that he had described many new Xylaria species, in

the early 1980's at one Florida foray many participants kept bringing him

specimens. Carris, a doctoral student at the University of Illinois at the time,

observed, “It was like they were bringing him gifts.... Even though he already

had a bagful of this fungus, he would look over each specimen very carefully

and say, ‘Oh, that is a beauty! I think I'll keep this one’ That speaks to who

he was and why people really liked him. He made everyone feel special.”

(JJanovich 2022)

‘Mr. XYLARIACEAE: Andrew Whalley, who contributed to an IMA

FuncGus celebration of Jack’s 80" birthday, confirmed that Jack’s doctoral

work “triggered a keen interest, and later a passion, for Xylariaceae. On

his appointment as Assistant Professor at the WSU faculty in 1963, with

positions in the Departments of Plant Pathology and Forestry and Range

Management, he continued with his studies of Xylariaceae, at first mainly on

chromosomes in Hypoxylon and later researching all aspects of the family.

In his remarkable and distinguished career, Jack, together with a number

of his many postgraduate students, established WSU as the world centre for

studies on Xylariaceae. He was also largely responsible for developing the

Mycological Herbarium at WSU, making it a renowned world-class facility

for educational and research purposes.

“Jack was President of the Mycological Society of America ... and his

Presidential Address “The Xylariaceae: systematic, biological, and evolutionary

aspects delivered at Tampa in 1977 during IMC2 demonstrated his clear

thoughts and knowledge of the family... If Julian Miller were alive today, he

would have been delighted and impressed with A REVISION OF THE GENUS

Hypoxy on by Yu-Ming Ju and Jack. Jack, Yu-Ming, and San Martin have

now revised 14 genera of the family and together with other former students

and colleagues published numerous papers on all aspects of Xylariaceae.”

“I am privileged to have known Jack since the mid-1970s, and to have

visited him and his family in Pullman ... and to have received them at our

home in North Wales. During his trips [to Wales] we visited the type locality

OCTOBER-DECEMBER 2021 ... XXXV

Fic. 2. Avid Fly Fisherman Rogers in 1982 proudly

displaying his 17-pound. steelhead, expertly §

rescued from Idahos Clearwater River.

~ Andrew Whalley

for Nemania chestersii in Anglesey, collected Hypoxylon rutilum in Llanbedr,

and Euepixylon udum by Aber Falls, Gwynedd, all rare species.... Considering

Pullman is hardly tropical, Jack has a broad and detailed knowledge of the

tropical species owing to his impressive global network of correspondents,

collaborators, and former students.... Jack has a wonderful sense of humour

and, together with his wife Belle, they have endeared themselves to all who

know them. He enjoyed hunting and fishing whenever he had the opportunity

and the 17 lb. Steelhead caught on the Clearwater River, Idaho, in November

1982 (FiG. 2) is true testimony to his success. (Whalley 2017)”

PaciFIC NORTHWEST FUNGI PRojECT (PNWEP). In 2002, Co-Directors Dr.

Joe Ammirati (University of Washington) and Dr. Dean Glawe (WSU Rogers

doctoral student and colleague) established a consortium of mycologists and

natural history enthusiasts cooperating to develop a comprehensive fungal

inventory of the Pacific Northwest (Alaska, British Columbia, Idaho, Montana,

Oregon, and Washington). Its mission was to facilitate cataloguing the fungi by

developing educational programs, databases, coordinating research projects,

and publishing an online journal initially under the name Paciric NORTHWEST

FunGI (Vols. 1-3(7); 2006-2008). Attending several meetings and forays, Jack

offered sage advice when needed. Our first meeting at Pack Forest near Mt.

Rainier in 2003 (Fic. 3) having convened shortly after his cataract operation,

Jack described in fascinating detail his laser surgery and the new lenses

slipped into place, rumbling with delight at being able to see once again in full

technicolor and—for the first time—sharply without glasses.

In mid-2008, the name of the journal was elevated to NorTH AMERICAN

Funa1 following Jack’s 80th birthday the previous year. The 267-page

FESTSCHRIFT IN HONOR OF PROFESSOR JACK D. ROGERS contained 17 papers

by 38 authors. Jumillera rogersii Y.M. Ju & H.-M. Hsieh and Camarops rogersii

Huhndorf & A.N. Mill. were among 14 taxonomic novelties; the many

XXXVI ... MYCOTAXON 136(4)

Fic. 3. Attendees of the first meeting of the PactrFIC NORTHWEST FUNGI PROJECT in Washington

State’s Pack Forest near Mt. Rainier National Park on 18 May 2003. Front (left > right): Michelle

Seidl, Lee Whitford, Lorelei Norvell, Lori Carris, Jennifer Falacy; 2‘” Row: John Dennis, Jacoba

Charles, Angela Muhlnickel, Claudia Nischwitz, Brenda Callan; 3"° Row: Gary Laursen, JACK

RoGers, Randy Bell, Joe Ammirati, John Hanna, Fred Rhoades; Back: Dennis Oliver, Bryce

Richardson, Ben Legler, Dean Glawe.

dedications touched on Jack's research philosophy (Watling & Milne: ‘in the

spirit of his understanding of how mycology should be conducted: to marry

classical morphological and ecological methods with modern techniques’),

heritage (Norvell & al.: to the indomitable West Virginian transplanted to

eastern Washington's Palouse’), friendship (Hemmes & Desjardin: ‘in fond

memory of wonderful collecting trips together in the Hawaiian Islands’), and

professorial mother-henning (Callan: ‘deep appreciation to Jack Rogers for

25 years of mentorship and friendship).

Marc Stadler

OCTOBER-DECEMBER 2021 ... XXXVII

Fic. 4. Jack “with the German amateur mycologists,

| Siegfried Woike (centre) and Hartmund Wollweber

(right) in the Neandertal near Wuppertal, Germany in

the summer of 2000, admiring a luxurious collection

of Daldinia concentrica.” [Stadler 2021]”

Marc Stadler first met Jack at the 1994 International Mycological Congress in

Vancouver, BC. “when we discussed my work on nematicidal activities of a

Daldinia sp. that I presented in a poster from my PhD thesis. We subsequently

started a collaboration that has laid the groundwork for my own taxonomic

research. I will never forget when he visited us in the Neandertal valley in

2000 and we showed him our local collection sites where we picked Daldinia

concentrica and other European Xylariales (Fic. 4).”

In his 2021 IMA FuNcus memoriam, Marc noted that once Jack focused

on the xylariaceous fungi, he became one of the foremost experts in the field.

“In retrospect, his monographs on important Xylariales ... appear even

more valuable. They helped [close] the gaps between classical ‘macromycete’

taxonomy and the molecular era via the establishment of holomorphic species

concepts. Jack dedicated several decades to the meticulous studies of both old

type material and freshly collected specimens in order to relate the taxonomic

concepts of the nineteenth century to the ‘modern’ era.” He and his students

did outstanding work on characterizing and cultivating the world’s Xylariales,

providing “numerous valuable descriptions that now allow other researchers

to establish asexual/sexual connections in this highly diverse and ecologically

versatile group.... Many of the phylogenetic relationships that Jack had

predicted from careful morphological studies in the 1980s and 1990s have been

confirmed in our recent work on polyphasic taxonomy, including multi-locus

phylogeny, chemotaxonomy, and phylogenomics.’ (Stadler 2021)

diodes WWW" =

ise of U Ye m\ JS

Fic. 5. Regents Professor Emeritus Jack D.

Rogers, “international expert on fungi,’

holds a microscopic slide stained with

Coccidioides-infected tissue from a dog

(Washington State Insider, 4 May 2014)

Shelly Hanks, WSU Photo Services

Bruce Andre

XXXVIII ... MYCOTAXON 136(4)

In addition to Jacks myriad professional achievements, he was also an

enthusiastic outdoorsman, a friend to many, and a legendary humorist. His

official obituary concludes: “He particularly enjoyed hunting pheasant and

quail on the breaks of the Snake River and fly fishing in rivers and lakes across

the Pacific Northwest. Jack was a regular at his two coffee groups as well as the

Pullman Presbyterian Church's Men's Bible Study and enjoyed visiting with his

many colleagues and friends. Jack loved a good joke or humorous story and

had the rare ability to remember and re-tell every entertaining anecdote he ever

heard. Whenever he said ‘that reminds me of a story..? we knew that whomever

he was with would soon be laughing until their sides ached.”

—Lorelei Norvell

Fic. 6. The Winter 2021 WasHINGTON

STATE INSIDER memoriam photo captures

Jack’s penetrating intelligence and loving

command of the forest, evoking the

conviction that any friend turning around

suddenly in the woods will find him

standing there.

LITERATURE CITED

Glawe DG, Ammirati JF. 2008. A Festschrift in Honor of Professor Jack D. Rogers. North American

Fungi 3(7): 1-4. https://doi.org/10.2509/naf2008.003.0071

Jovanich A. 2022 (“2021”). In memoriam: Jack Rogers. Washington State Magazine. Winter 2021:

https://magazine.wsu.edu/2021/11/08/in-memoriam-jack-rogers/

Ju YM, Rogers DJ. 1996. A revision of the genus Hypoxylon. Mycologia Memoir no. 20. APS Press,

St. Paul. 365 p.

Rogers JD, Callan BE, Samuels GJ. 1987. The Xylariaceae of the rain forests of north Sulawesi

(Indonesia). Mycotaxon 29: 113-172.

Stadler M. 2021. Jack D. Rogers (1937-2021), in DL Hawksworth (ed.), MycoNews. IMA Fungus

12: 36. https://doi.org/10.1186/s43008-021-00085-9

Whalley AJ. 2017. Jack Rogers—Mr Xylariaceae turning 80. IMA Fungus 8:18-19 [Whalley name

omitted in original article, cited as Hawksworth DL, Seifert KA, Senn-Irlet B & al. Awards and

Personalia. IMA Fungus 8: A16—A24]. https://doi.org/10.1007/BF03449430

OcCTOBER-DECEMBER 2021 ... XXXIX

IN REMEMBRANCE—

KALMAN VANky (1930-2021)

Fic. 1. KALMAN VANKy. Undated photo of

the author scanned from the dedication

page of EUROPEAN SMUT FUNGI (1994).

Dr. KALMAN GEzA VANKyY was borna Szekely- Hungarian in Székelyudvarhely

(Odorheiu Secuiesc, colloquially Odorhei), Transylvania (Romania), on

15 June 1930. His website (accessed in February 2022) reflects his restless

nature, citing Kalman as the married father of three with the same German

Tubingen address I visited in 2005, a Swedish email address signifying his

2014 move to that country, and joint Hungarian and Swedish citizenships.

Colorful, indefatigable, enthusiastic, irrepressible, and productive are all

terms that friends and colleagues use to refer to this world authority on smut

fungi, who died on 12 October 2021 at the age of 91.

After earning his BS and MS in Biology from the University of Bucharest

(Romania) in 1953, Kalman spent the next four years working as a researcher

for the Department of Phytopathology of Bucharest’s Agricultural Research

Institute. After earning his MD in 1961 from the University of Medicine and

Pharmacy in Tirgu Mures, he practiced medicine in Odorhei and Borsec (a

Romanian resort known for its mineral water springs), becoming intrigued

by plant diseases. His interest in smut fungi (Ustilaginomycotina) having been

piqued by the lectures of Prof. Traian Savulescu, founder of the Romanian

School of Phytopathology and president of the Romanian Academy, Kalman

XL ... MYCOTAXON 136(4)

Fic. 2. LEFT: Undated early photo of Kalman at the microscope. MIDDLE: Kalman at age 60 (1991).

RIGHT: Prof. Terence Ingold and Kalman talk during IMC IV in Regensburg (1990).

began collecting “as many different species as possible,’ a hobby that soon grew

into his life’s work.

In 1969, Kalman moved to Gagnef, Sweden, where he continued working

as a family physician, but only half-time; during 1970-86 he also worked as

a “self-financed, half-time” smut fungus taxonomist, eventually securing a

scholarship to Uppsala University, where he earned his PhD in Botany under

the supervision of Prof. Johan Axel Nannfeldt, an expert on pathogenic rust

fungi, smut fungi, and Exobasidium. Kalman successfully defended his thesis

on Carpathian Ustilaginales in 1985.

In 1986 he moved to Germany to focus exclusively on smut fungi, working

as a taxonomic researcher for Prof. Franz Oberwinkler in the Department

of Special Botany and Mycology at the University of Tubingen, maintaining

his usual frenetic work pace: “collecting, identifying, describing, illustrating,

classifying, and publishing species and genera from all over the world.” By his

1995 ‘retirement, he had already collected ustilaginomycetes in Europe, Africa,

Asia, Australasia, and Central & North America, but his pace actually increased

as he worked independently from his homes in Germany and Sweden. By his

sixth collecting trip to Australia in 2017, the peripatetic Kalman had added

South America to his list, having (by his count) collected in 56 countries.

[His website, however, missed Canada when he catalogued British Columbia's

Victoria as part of the United States, and a few ‘countries’ represent unique

regions rather than political entities. ]

HERBARIUM USTILAGINALES VANKY (HUV)—Having begun a private

herbarium in Romania in 1954, Kalman continued to add specimens from all

over the world while living in Sweden and Germany. By January 2013, HUV

OCTOBER-DECEMBER 2021 ... XLI

yo ae

So sii 2

Fic. 3. LEFT: Kalman collecting in Costa Rica in 1991. RIGHT: Kalman & Christine look on as

Tilletia co-author Prof. N.D. Sharma inspects a specimen during their 1992 trip through India.

contained 22,050 collections, including a great number of types (holotypes,

isotypes, neotypes, lectotypes, syntypes, paratypes, topotypes), apparently

including ‘portions of types from other collections, an issue that occasioned

considerable controversy’ (Hawksworth 2021).

Kalman exchanged smut fungi for many years on a one-to-one basis and

distributed exsiccata numbers 1-1350 comprising 76,500 samples from ‘Vanky,

Ustilaginales exsiccata (1960-2010). The exsiccata contained smut fungi from

all over the world, and Vanky was justly proud that it represented “the sole

exsiccata of smut fungi appearing after World War II” Approximately 76,000

specimens (Nos. 1-800 with 70 fascicles, Nos. 801-1000 with 45 fascicles, and

Nos. 1001-1350 with 33 fascicles) were distributed to mycological herbaria or

sent to mycologists from all over the world. The exsiccata represented 615 smut

fungus species (including 164 types) and 68 genera on 825 host plant taxa.

In October 2013, HUV was relocated to the Queensland Plant Pathology

Herbarium (BRIP) in Brisbane, Australia, where it is now curated by Dr. Roger

G. Shivas, Kalman’s co-author, fellow collector, and friend. On announcing

the relocation, David Hawksworth (2013) noted, “It is gratifying that this

important collection is now safeguarded for future generations, and I am sure

all mycologists will wish Kalman and his wife many relatively smut-free and

more relaxing years!”

PUBLICATIONS—Kalman was 26 when he published (as co-author) his first

paper on the pathogenic gram-negative bacterium that caused variety of plant

diseases, including ‘black rot’ in crucifers, Xanthomonas campestris (Bucur &

Vanky 1956). Seven years later, he had shifted his focus to the smut fungi that

he would eventually master (Vanky 1963). He named his first taxon in 1975

XLII ... MYCOTAXON 136(4)

Fic. 4. LEFT: Kalman collecting on the equator in Ecuador (1993).

RIGHT: Kalman searching for possibly smut-infected hosts in South Arica (1996).

(Vanky 1975: Doassansia alismatis-oligococci) and co-authored his final two

taxa forty years later (Riess & al. 2015: Talbotiomyceae, Talbotiomycetales).

Kalman and his co-authors named a total of 13 families, 46 genera, and at

least 410 new species, also publishing 19 noms. nov. and making 449 new

combinations.

In total, Kalman published ten substantial and definitive treatises on

the Ustilaginales, the three most comprehensive of which were his 570-page

EUROPEAN SMUT FUNGI (1994), 1458-page SMuT FUNGI OF THE WoRLD (2012),

and the third 288-page edition of ILLUSTRATED GENERA OF SMUT FUNGI

(2013). His critically acclaamed monographs, which supplanted Zundel’s 1953

THE USTILAGINALES OF THE WoRLD as the ultimate world authority on smuts

were all lavishly illustrated, containing “detailed nomenclators, descriptions

drawings, photographs, SEM micrographs, and host lists for the world’s 1650

known smut species.” (Hawksworth 2021).

In addition to his books, Kalman published 224 papers, 56 of which were

published in Mycotaxon. My first encounter with this major mycological

force involved his “Taxonomic studies on Ustilaginomycetes 24’ (Vanky 2004),

which I received immediately after being appointed Editor-in-Chief. Kalman

had published in the journal since 1982 and was well-trained in our hard-

copy photo-ready procedure; the manuscript was obviously in impeccable

shape, leaving little for me to do but approve, paste on masthead and page

numbers, send to the press, and mail the drawings and photos back to

Germany after publication.

OCTOBER-DECEMBER 2021 ... XLIII

ted

Fic. 5. LEFT: Eric H.C. McKenzie and Kalman collecting in New Zealand (1998).

RIGHT: Kalman standing next to a large termite mound in Uganda (2002).

BOTTOM: Son Tomas Vanky with proud father on a 2003 collecting trip through Mexico.

After the 2004 change from postal to electronic submission accompanied

by style and formatting changes, followed by Shaun's addition as Nomenclature

Editor in 2005, however, things became more interestingly complex, launching

an era of increased editorial-author email exchange and ‘artful’ compromise.

Kalman’s copy, prepared beautifully by wife Christine (who only later admitted

that our innovations presented a real challenge), now needed numerous

editorial adjustments, and the author tended to question each and every change.

XLIV ... MYCOTAXON 136(4)

\\\ eee TT a

{ |e amen “ 3

Fic. 6. LEFT: Gracious hosts Kalman, Christine (and Bernese Mountain Dog Tara looking for

treats) lead newlyweds Lois and David Lehwalder on an evening stroll of Tubingen in 2005.

TOP RIGHT: Christine and Kalman cleaning the kitchen before a late-night exchange of

mycological banter with visiting MycoTaxon EIC Norvell. BoTTOM RIGHT: Undated photo of

Herbarium Ustilaginales Vanky (HUV).

Christine's customary hard returns and page breaks wrought considerable

havoc in Pagemaker® and InDesign®, necessitating considerable negotiation

during a rather steep learning curve for both sides. Fortunately, we emerged

victorious and friends, with my library shelves rewarded by a lovely copy of

EUROPEAN SMUT FUNGI.

Learning that I would be driving with my father and brand-new stepmother

through Germany to attend the 2005 International Botanical Congress in

Vienna, Kalman insisted we stop in Tiibingen. July 9 found us enjoying

Christine's elderberry wine, hors doeuvres ‘sans sel’ (which an eager Kalman

served too soon), and delicious dinner and then striding at dusk past lighted

shop windows led by a ‘wild’ Hungarian, Christine, and Bernese Mountain Dog

Tara along the Neckar River and over cobblestones among Tubingen students

and residents. After dropping the ‘newlyweds’ off at the Hotel Krone, Kalman

and I discussed mycology and journals until well after midnight before I retired

to my comfortable bed in the living room next to HUV. Our Tubingen stay

was magical and voted unanimously the highpoint of our European trip. After

our return, I received a note telling me that my contact lens solution had been

left in the bathroom. “Should we send it to you? ...We prefer to keep it until

you will come again to us, that time for a longer period. It was really a nice

but too short stop on your way from Frankfurt to Vienna. Maybe next time it

Left & Top right: Lorelei Norvell

OCTOBER-DECEMBER 2021 ... XLV

Fic. 7. 2005. LEFT: Fellow Roger G. Shivas and Kalman collecting near Cairns in Queensland,

Australia. RIGHT: Kalman, Swiss rust expert Reinhard Berndt, and Christine chat during the final

social banquet at the International Mycological Congress in Cairns.

will be longer? I have much to do, especially with the preparation of the world

monograph. Therefore, I am ending this too-short letter. Christine and I send

you our warmest regards and Tara her nicest tail-wagging.”

Their travels did not always go smoothly. Recapitulating their 2006 travels

through Hungary and Romania after visiting relatives and friends, Christine

wrote, “Kalman had the roots of three teeth removed and two ‘new teeth

implanted. All in one session! In the evening after that, he had a proper

dinner—but one that did not need too much chewing. After that exhausting

experience, we wanted to spend a few days of quiet collecting in Slovenia. The

first day we had pouring rain and storm, the second day Tara became seriously

ill with high fever, neither wanting to get up nor eat a single bite. We hurried

home to see our vet who finally found out she has Babesiosis and probably

Ehrlichiosis, both transmitted by the same hungry Hungarian tick! She feels

much better now, but the results of her blood tests are still very bad. Kalman is

looking forward to see the PDF file of the manuscript and we both are looking

forward to seeing you in Cairns’’

We did meet in Cairns during the 2005 International Mycological Congress

and had an especially convivial time at the closing reception in the “Tanks, but

Kalman wrote about their ‘rather adventurous’ return, “We missed our flight

in Cairns and [spent] long waiting times in several airports [before flying]

Cairns—Sydney-Singapore-London-Stuttgart. Finally, tired after over 36

hours, we arrived home without our luggage, which showed up two days later

in bad condition. Now, we are planning our next collecting trip to Cameroon

[February/March 2006] in connection with the 18th AETFAT Congress, where

I will present a talk about smut fungi of Africa.”

XLVI ... MYCOTAXON 136(4)

(2012).

’

' > . r r r

{ iL ) 7 Fic. 8. A photogenic Kalman Vanky at age 82

: ; -

After publishing his last paper with us in 2009, Kalman wrote that he would

be finishing his mycological endeavors. “At the moment my main goal is to co-

operate with the APS Press in publishing the World Monograph which could

take up to one year from now. I would like also to assure that my collection will

be deposited in a good herbarium and preserved under optimal conditions for

coming generations.... Finally, I would like to express once more my thanks and

gratitude for publishing my manuscripts in MycoTaxon, a journal which you

succeeded within a few years to transform into a journal of highest scientific

quality and aesthetic level. [I strongly suspect that you have in your vessels also

the blood of a magician]. With all best wishes, Kalman.”

AWARDS AND HONORS: Kalman, who became a proud honorary member

of the Hungarian Academy of Science in 2001, also received the Pazmany

Dénes Award & Laudatio from the Kalman Laszl6 Mycological Society of

Transylvania (Romania) (2012), the Arany Janos-Medal of the Hungarian

Academy of Science (2014), and the Officer’s Cross of the Hungarian Order

of Merit (2020). Three species (Anthracoidea vankyi, Orphanomyces vankyi,

Uromyces vankyorum) and three genera (Kalmanomyza, Kalmanago, and

Vankya) have been named in his honor.

Kalman’s ashes were to be interred in Vajdahunyadalso (Romania) beside his

first wife, Dr. Magda Vanky (1936-1970), also a physician. Surviving Kalman

are Christine; sons and daughter Dr. Kalman Vanky Jr. Dr. Tomas Vanky,

OCTOBER-DECEMBER 2021 ... XLVII

Dr. Elisabet Vanky; 10 grandchildren; numerous relatives, mycological

colleagues, and friends; and his last wishes in the form of a quote from the

Hungarian poet (partly translated by Kalman himself):

I have done all I could,

I have paid those I thought I should.

I am releasing everyone's tribute and, please,

Forget the declining earthly copy of my face.

Kalman, we regret we must use our editorial license to overrule your wishes

one last time.

—Lorelei Norvell

LITERATURE CITED—

Bucur E, Vanky K. 1956. Contributii la studiul putregaiului bacterian al verzei

[Xanthomonas campestris (Pammel) Dows.]. - Comun. Acad. Republ. Populare

Romane, St. Agricole 6: 1111-1115.

Hawksworth DL. 2021. MycoNews. IMA Fungus 12: 36.

https://doi.org/10.1186/s43008-021-00085-9

Hawksworth DL, Luangsa-ard JJD, Crous PW & al. 2013. News. IMA Fungus 4,

A35-A38. https://doi.org/10.1007/BF03449309

Riess K+, Bauer R, Kellner R, Kemler M, Piatek M, Vanky K, Begerow D. 2015.

Identification of a new order of root-colonising fungi in the Entorrhizomycota:

Talbotiomycetales ord. nov. on eudicotyledons. IMA Fungus 6: 129-133.

https://doi.org/10.5598/imafungus.2015.06.01.07

Vanky K. 1963. Citeva specii noi de Ustilaginale pentru micoflora R.P.R. - Lucr. Grad.

Bot. Bucuresti. Acta Botanica Horti Bucurestiensis 1961-—62:201-211.

Vanky K. 1975. Doassansia alismatis—oligococci Vanky, sp. nov. — Svensk Bot. Tidskr.

69: 45-48.

Major VANKY PUBLICATIONS—

Vanky K. 1985. Carpathian Ustilaginales, Symbolae Botanicae. 309 p.

Vanky K. 1994. European smut fungi, Fischer Gustav Verlag GmbH & Co. KG 570 p.

Vanky K. 2007. Smut Fungi of the Indian Subcontinent, Polish Botany Institute, 265 p.

Vanky K. 2012. Smut Fungi of the World. APS Press, St. Paul. 1458 p.

Vanky K. 2013. Illustrated Genera of Smut Fungi (3 ed.) APS Press, St. Paul. 288 p.

Vanky K, McKenzie EHC. 2002. Smut Fungi of New Zealand (6 ed.), Fungal Diversity

Press, 259 p.

Vanky K, Oberwinkler F. 1994. Ustilaginales on Polygonaceae: a taxonomic revision.

Nova Hedwigia. 96 p.

Vanky K, Shivas RG. 2007. Fungi of Australia: The Smut Fungi, Melbourne, Australian

Biological Resources Study Canberra and CSIRO Publishing, 267 p.

Unless otherwise noted, all facts and photographs above were obtained from Kalman’s still active

website: www.kalman-vanky.de/index.html.

MY COTAXON

October-December 2021—Volume 136, pp. 693-717

https://doi.org/10.5248/136.#693

Lamprospora benkertii sp. nov., and an evaluation of

Lamprospora spp. with seaveri-type ascospore ornamentation

JAN ECKSTEIN'S, MARCEL VEGA’,

ZUZANA SOCHOROVA®®, LUKAS JANOSIK*S

' Arnoldiweg 20, Gottingen 37083, Germany

? Kohlhoefen 17, Hamburg 20355, Germany

> Department of Botany, Faculty of Science, Palacky University Olomouc,

Slechtitelii 27, Olomouc 783 71, Czech Republic

* Department of Botany, Faculty of Science, Charles University,

Bendtska 2, Prague 128 01, Czech Republic

CORRESPONDENCE TO: ' jan.eckstein@octospora.de

° asco.sochorova@gmail.com * janosiklu@natur.cuni.cz

ABsTRACT—Lamprospora benkertii is described as a new species based on collections

from Georgia, Germany, Norway, Slovakia, and Switzerland. It is characterized by

orange apothecia without a fimbriate margin, globose ascospores with a seaveri-type

ornamentation comprising strong ridges forming a reticulum anda secondary reticulum

formed by fine ridges within the meshes of the primary reticulum. The fungus infects

rhizoids of Trichostomum crispulum. Delineation of the new species is supported by LSU

and EFla sequence analyses. Lamprospora benkertii is compared with other Lamprospora

species with a seaveri-type or similar ascospore ornamentation: L. ascoboloides, L. cailletii,

L. dicranellae, L. funigera, L. hispanica, L. irregulariata, L. leptodictya, L. norvegica,

L. paechnatzii, L. seaveri. We also present an updated key for this group of species as

well as a more complete description of L. hispanica based on our observations from

several collections. New host specificity data are provided for L. hispanica, L. leptodictya,

L. norvegica, and L. paechnatzii.

Key worps—bryophilous Pezizales, cryptic diversity, host specificity, integrated taxonomy,

Pyronemataceae

SORCID identification numbers [0000-000...]: ECKSTEIN: 2-7214-3321; VEGA: 2-0176-9503;

SOCHOROVA: 3-0022-9780; JANOSIK: 2-4001-0145.

694 ... Eckstein & al.

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Lamprospora benkertii sp. nov. (Europe) ... 695

Introduction

Lamprospora De Not. exhibits a remarkable variability in ascospore

ornamentation. Because it may consist of variously shaped ridges, warts,

spines, tubercles, and bands, both alone and in combination, ascospore

ornamentation is traditionally considered the most important character for

species delimitation. In a monographic work, Benkert (1987) distinguished six

basic types of ornamentation in Lamprospora. One type, characterized by long

curved ridges, sometimes forming loops and often tapering at interjections or

ends, was termed by him as seaveri-type (referring to L. seaveri).

In recent years, integrated taxonomy combining traditional morphological

studies with new molecular techniques has proved a powerful tool in assessing

biodiversity in all groups of organisms (e.g., Caparrds & al. 2016, Deniz & al.

2015, Lépez-Bautista & al. 2006, Nater & al. 2017, Nedeljkovic & al. 2015, Zhao

& al. 2019). Only recently has this approach also been applied to bryophilous

Pezizales (Vega & al. 2017, 2019, Sochorova & al. 2019). One of the main

discoveries from molecular investigations of bryophilous Pezizales was the very

high host specificity of all investigated species. In consequence, host identity

is even more important for delineating bryophilous Pezizales species than

ascospore ornamentation (Egertova & al. 2018). This makes species reported

from multiple, not closely related, hosts likely candidates for hidden diversity

and due for evaluation.

Eckstein (2014) reported Lamprospora hispanica, one species with seaveri-

type ascospore ornamentation, as new to Germany, citing collections on

Aloina sp. and on the hitherto unknown host Trichostomum crispulum Bruch

(Pottiaceae). Since then, we studied many more collections on both hosts and

clarified that two species with similar ascospores were at hand. The aim of

the present article is to describe the new species on Trichostomum crispulum

and to compare it with other Lamprospora species with seaveri-type ascospore

ornamentation.

Material and methods

Sample collection and observation

The description of macroscopic and microscopic characters is based on vital

material. Microscopic structures were measured in tap water using standard light

microscopy. For each collection, at least 20 ascospores and 10 other structures were

measured. In all species, ascospore dimensions always include ornamentation, unless

stated otherwise. Ascospore ornamentation was studied in cotton blue, and non-

amyloidity checked in Lugol’s solution. Scanning electron micrographs (SEM) were

taken from air-dried samples.

696 ... Eckstein & al.

Herbarium acronyms follow Thiers (2019). Lists of collections are ordered

alphabetically by country and from north to south within countries. Collector

abbreviations include CN (Csaba Németh), DB (Dieter Benkert), EB (Emiel

Brouwer), EP (Eckehard Paechnatz), ES (Elisabeth St6ckli), GE (Gtinter Eckstein),

GM (Gianfranco Medardi), HD (Henry Dissing), JE (Jan Eckstein), LJ (Lukas

Janosik), MdIT (Margarita de la Torre), MS (Michal Sochor), MV (Marcel Vega), RK

(Roy Kristiansen), RMG (Rubén Martinez-Gil), RTL (Raul Tena Lahoz), SS (Sigmund

Sivertsen), TR (Torsten Richter), ZE (Zuzana Egertova).

DNA extraction, PCR amplification, sequencing

DNA was extracted from fresh, dried, or CTAB-stored apothecia using the CTAB

method (Doyle & Doyle 1987) or the Zymo Research Fungal/Bacterial kit. Amplification

was carried out for the 28S subunit of ribosomal DNA (LSU) using primers LROR

and LR6 (Vilgalys & Hester 1990) and for the translation elongation factor-lalpha

(EFla) with primers EF1-983F and EF1-1567R (Rehner & Buckley 2005). PCR was

performed with Kapa polymerase, following a standard protocol with 37 cycles and

annealing temperature of 56°C. The PCR products were purified by precipitation with

polyethylene glycol (10% PEG 6000 and 1.25M NaCl in the precipitation mixture)

and sequenced in both directions using the Sanger method (Macrogen Europe, The

Netherlands and Sequencing laboratory of the Faculty of Science, Charles University,

Prague, Czech Republic).

Phylogenetic analysis

LSU rDNA sequences analyzed included nine sequences of L. benkertii, eight

of L. hispanica, one or two of other Lamprospora species with similar ascospore

ornamentation, and Octospora leucoloma Hedw. as outgroup. Additionally, EFla was

analyzed in selected collections of L. benkertii and L. hispanica. Along with sequences

newly generated in this study, relevant data from the GenBank database were used

(https://blast.ncbi.nlm.nih.gov; sequences originate from Perry & al. 2007, Hansen &

al. 2013, Vega & al. 2017, and Egertova & al. 2018). Sequences were assembled, edited,

and aligned using GENEIOUS (ver. 7.1.7., Biomatters). Bayesian phylogeny inference

(BI) was computed in MrBayeEs (ver. 3.2.4, Ronquist & al. 2012) using the GTR+I+G

substitution model, as determined by AICc in JMODELTEST (ver. 2.1.4, Darriba &

al. 2012). The analysis was run for 10 million generations in two independent runs,

sampling every 1000th generation and excluding the first 25% of generations as burn-

in. Sequences and GenBank accession numbers used in the phylogenetic analysis are

listed in TABLE 1.

Taxonomy

Lamprospora benkertii Eckstein, M. Vega, Sochorova & Janogik, sp. nov. Fics 1-4

MB 831135

Differs from similar Lamprospora species by its apothecia without a fimbriate margin,

globose ascospores with an alveolate reticulum formed by strong ridges, and infections

located on rhizoids of Trichostomum crispulum.

Lamprospora benkertii sp. nov. (Europe) ... 697

Fic. 1. Lamprospora benkertii (A, C. holotype; B. B 70 0100991).

Apothecia on soil between shoots of Trichostomum crispulum.

Type: Germany, Saxony-Anhalt, Western slope of Spielberger Hohe 1 km ENE Spielberg,

51.3247°N 11.5992°E, 220 m a.s.l, on loamy-calcareous soil with Trichostomum

crispulum, Didymodon fallax, and Ephemerum recurvifolium, 12.VIII.2011, JE22084

(Holotype, B 70 0100994).

Etymo tocy: honoring Dieter Benkert for his meritorious service in the taxonomy of

bryophilous Pezizales

APOTHECIA gregarious on soil between shoots of Trichostomum, (0.2-)0.4-1

(-1.5) mm in diameter, first spherical, becoming cup-shaped, finally discoid

with a flat hymenium, sessile, with few hyaline and flexuous anchoring hyphae

at the base; MARGIN inconspicuous; HYMENIUM orange to bright orange, outer

surface slightly paler.

698 ... Eckstein & al.

Ascr 180-250(-325) x 18-28 um, cylindrical to slightly club-shaped,

8-spored, unitunicate, operculate, inamyloid, arising from croziers.

ASCOSPORES uniseriate, hyaline, thin-walled, globose, 13-16.5 um in

diameter (ornamentation included) or 12-14.5 um without ornamentation,

always with a lipid drop, 8-10(-11) um diameter; ORNAMENTATION of the

seaveri-type consisting of curved ridges approximately 0.8-1.2(-1.6) um wide

and 0.8-1.2 um high, forming a coarse alveolate reticulum with 4-8 meshes

across ascospore diameter, meshes fairly variable and always with a thin

secondary net of fine ridges, ridges rarely with loops, sometimes circling the

ascospore and rarely ending blind, anastomosing ridges often tapered towards

intersection, ornamentation cyanophilous, quickly dissolving in potassium

hydroxide (10% KOH).

ParAPHYSES filiform, straight, pluriseptate, rarely forked and only in lower

third, gradually widening from 2.5-3 um in lower third to 4-6(-8) um at tip,

terminal cell (30-)45-85(-100) um long, with distinctive carotenoid pigment

turning cyan to olivaceous in Lugol's solution, and numerous vacuolar bodies,

size <4 um.

SUBHYMENIUM consisting of thin-walled, variously shaped, hyaline cells

2.5-6 um broad, merged with the medullary excipulum; MEDULLARY

EXCIPULUM about 60-90 um thick, comprising thin-walled, variously shaped

cells forming a textura intricata; ECTAL EXCIPULUM ca. 40-55 um thick,

comprising thick-walled globose, subglobose to subangular cells (textura

subglobulosa), 6-20 x 5-18 um, cell wall <3 um thick; Marco of parallel

hyphae with thick-walled end cells 20-50 x 4-10 um not (or minimally)

protruding over the hymenium (textura prismatica-porrecta).

INFECTIONS located on rhizoids of Trichostomum crispulum, consisting of

mostly 1-celled ellipsoid appressoria, 20-30 x 10-12 um, sometimes with a

somewhat enlarged adjacent cell, initially free, becoming completely covered

with anchoring hyphae; INFECTION PEG ca. 1 um wide, surrounded by a tube

of rhizoid cell-wall material, forming intracellular haustoria; PENETRATION

POINT surrounded by a circular thickening of the rhizoid wall; HausToRta often

ramified with irregular shape and without cross walls, not growing through

rhizoid walls; GROWTH MODIFICATIONS with inflated cells and more cross walls

Fic. 2. Lamprospora benkertii (A, F, H. holotype; B. B 70 0100987; C. PRC 4580; D, E, G. PRC

4601). A-C. Ascospores stained with cotton blue. D. Ascospores and paraphyses in water.

E. Upper part of paraphyses in water. F Marginal hyphae of an apothecium seen from outside

Lamprospora benkertii sp. nov. (Europe) ... 699

stained with cotton blue. G. Cross section of a vital apothecium showing margin, excipulum,

and hymenium. H. Cross section of a dried apothecium showing subhymenium, medullary,

and ectal excipulum. Scale bars = 20 um.

700 ... Eckstein & al.

Fic. 3. Lamprospora benkertii (A-B. holotype; C-D. B 70 0100987).

Ascospores (SEM). Scale bars: A-C. = 10 um; D. = 20 um.

than normal sometimes observed on infected and adjacent rhizoid cells; HosT

not weakened discernibly by the infection.

ECOLOGY & DISTRIBUTION—On soil in open situations like grassland, or

along tracks and paths together with bryophytes including, apart from the

host, Aloina spp., Didymodon fallax, D. acutus, Ephemerum recurvifolium,

and Weissia longifolia. Known from Georgia, Germany, Norway, Slovakia, and

Switzerland.

ADDITIONAL SPECIMENS EXAMINED—GEORGIA. RACHA-LECHKHUMI AND KVEMO

SVANETI, Racha mountains, 5 km E of Shovi, 42.6911°N 43.7317°E, 2240 m a.s.l.,

27.V1.2014 ZE (PRM 952408). GERMANY. Saxony-ANHALT, NE Grockstadt,

51.3331°N 11.5931E, 220 m a.s.l., on calcareous soil in dry grassland, 21.VIII.2017

JE51316 (B 70 0100984); 2.35 km NNE Reinsdorf, 51.3128°N 11.6183°E, 195 m

a.s.l., on south-facing slopes of loess-soil with Trichostomum crispulum, 11.X.2011

JE22454 (B 70 0100996). NORWAY. NorDLAND, Saltdal, Junkerdalen, Gammelveien

innenf. bommen, 120 m a.s.l., between mosses on track, 31.VIII.1988 SS88-78 (TRH

F-10783 as Lamprospora cf. ascoboloides); 29.VIII.1998 HD&SS98-14 (TRH F-10639 as

Lamprospora ascoboloides). SLOVAKIA. ZILINA, Sulov-Hradna, 49.1576°N 18.5808°E,

425 ma.s.L, margin of a forest way, 20.VIII.2016, ZE (PRM 952409); TRENCIN, Tren¢in,

48.8876°N 18.0610°E, 220 m a.s.l., between mosses on the margin of a small forest path,

Lamprospora benkertii sp. nov. (Europe) ... 701

Fic. 4. Lamprospora benkertii (holotype). Several infectious structures in optical section on

rhizoids of Trichostomum crispulum; fungal cells dotted. Scale bar = 20 um.

10.XII.2014 LJ (PRC 4579); Beckovské Skalice, 48.7744°N 17.8944°E, 220 m a.s.l., on

path in grassland, 07.X.2014 LJ (PRC 4580); 27.X.2019 LJ (PRC 4601). SWITZERLAND.

BERN, Gasteretal, 46.4672°N 7.6611°E, 1350 m a.s.l., on soil, 30.I[X.2016 ES (B 70

0100991); Kandersteg, 46.4744°N 7.6572°E, 1240 m a.s.l., on soil in grazing land,

25.VUI.2018 MV180825-2 (B 70 0100987); 46.4447°N 7.7225°E, 1580 ma.s.l., on soil in

grazing land, 04.X.2013 ES (B 70 0100982).

PHYLOGENETIC ANALYsIs—Lamprospora benkertii forms a well-supported

clade with very little polymorphism among individual collections (Fic 5).

In LSU, five collections are fully identical (MN394601 - holotype, MN400066,

MN394592, MN394614, and MN394602), with three others differing only in

a single nucleotide (MG949136, MN394603 and KC012684 — published as

L. norvegica by Hansen & al. 2013) and one other in another single nucleotide

(MN394594). Lamprospora paechnatzii was identified as the closest relative of

L. benkertii; L. hispanica, which is morphologically very similar to L. benkertii,

differs strongly both in LSU and EF 1a (data not shown). A similar phenomenon

was also observed in L. seaveri sensu lato, which was described as associated

with Ceratodon purpureus and Bryum sp. (Benkert 1987); however, collections

from these two hosts form independent lineages.

702 ...

Eckstein & al.

Lamprospora hispanica MN400064

Lamprospora hispanica MN394593

Lamprospora hispanica MN394598

Lamprospora hispanica MN394599

Lamprospora hispanica MN394596

Lamprospora hispanica MN394600

Lamprospora hispanica MN394607

Lamprospora hispanica MN394608

0.92° Lamprospora aff. seaveri MN394609

Lamprospora aff. seaveri MN394595

Lamprospora ascoboloides DQ220358

Lamprospora sp. DQ220361

Lamprospora dicranellae KY858952

Lamprospora dicranellae KY858955

Lamprospora leptodictya MN394610

Lamprospora leptodictya MN394611

Lamprospora seaveri MN394612

Lamprospora seaveri MN394597

Lamprospora norvegica MN400065

Lamprospora norvegica MN394606

Lamprospora cailletii MN394604

Lamprospora cailletii MN394605

Lamprospora paechnatzii MN394613

Lamprospora benke:

Lamprospora benkertii| M

Lamprospora benkertii M

Lamprospora benkertii MN39

Lamprospora benkertii M

Lamprospora benke:

Octospora leucoloma MK569370

Fic. 5. Bayesian phylogram of Lamprospora spp. based on LSU rRNA sequences, with Octospora

leucoloma as outgroup. Bayesian posterior probabilities are shown above branches.

ComMENTS— Trichostomum crispulum, the host of the new L. benkertii, is not

known as a host of other bryophilous Pezizales. It has a circumpolar southern-

temperate distribution with occurrences in Europe, Asia Minor, Siberia, East

Asia, New Guinea, Macaronesia, North and Central Africa, Newfoundland,

and Central America (Smith 2004).

In his main contribution to the taxonomy of Lamprospora, Benkert (1987)

listed six species with seaveri-type ascospore ornamentation: L. cailletii,

L. hispanica and L. seaveri, as well as L. ascoboloides, L. leptodictya and

L. paechnatzii. The first three species were assigned to the subtype ‘seaveri,’

characterized by anastomosing ridges forming an alveolate reticulum, and the

latter three to the subtype ‘paechnatzii, delineated by only partly anastomosing

ridges not forming a reticulum. Subsequently, another species, L. norvegica,

was also described with seaveri-type ornamentation (Benkert & al. 1991).

Two little known species, L. irregulariata from India and L. funigera from

Australia, also must be considered in this context. Additionally, we include

L. dicranellae Benkert in this discussion, because in the past it has frequently

been mistaken for L. ascoboloides. We discuss these species in alphabetical

order below with delineations based on our own observations except for L.

funigera and L. irregulariata.

Lamprospora benkertii sp. nov. (Europe) ... 703

Lamprospora ascoboloides Seaver, Mycologia 6: 10. 1914.

Based on collections from the US states Connecticut, New York, and Virginia,

L. ascoboloides is characterized by globose ascospores 13-15(-17) um in

diameter [12-14 um in Benkert (1987) and (14—)16-18 um incl. ornamentation

in Wang & Kimbrough (1992)], ornamented by curved ridges 1-2 um wide

and 1-1.5(-2) um high that often end bluntly and do not (or hardly) form a

rudimentary reticulum (Seaver 1914, Benkert 1987, Wang & Kimbrough 1992).

Furthermore, the ornamentation lacks warts. Our study of the NY type material

revealed a dicranelloid moss as the host. (probably Dicranella heteromalla, but

with juvenile shoots too small for certain identification). Notably, the infection

induces spherical galls 40-80 um in diameter on the rhizoids. These rhizoid

galls are completely covered with a layer of hyphae. L. ascoboloides differs from

L. benkertii in broader ridges and the host.

The name L. ascoboloides was later applied many times to collections with

strong, curved ridges as ascospore ornamentation, but critical studies of these

specimens always led to other identifications. For instance, L. ascoboloides

in reports by Caillet & Moyne (1980) was later re-determined by Benkert

(1987) as L. dicranellae. The description and illustrations of “L. ascoboloides’

by Schumacher (1993) also clearly show L. dicranellae. Our revision of

L. ascoboloides specimens in TRH indicated mostly L. dicranellae or rarely

L. arvensis (Velen.) Svréek as well as several yet undescribed species. We

also revised a specimen from K (listed in Medardi 2006) to L. dicranellae.

We also studied two collections sequenced by Perry & al. (2007, GenBank

accession numbers DQ220358 and DQ220361) that also have L. dicranellae-

like ascospores, but their sequences do not cluster with other L. dicranellae

samples (Vega & al. 2017), clearly indicating that further study is needed.

Dicranella species were not found as hosts in any of these cases. Medardi

(2005) reported L. ascoboloides from Italy accompanied by Dicranella,

Ditrichum, Riccia, and Anthoceros. Unfortunately, our investigation of MCVE

15993 resulted in no unambiguous observation of the infection but the host is

most likely Dicranella or Ditrichum. Therefore, the specimen could represent

L. ascoboloides but without definite identification of the host, its identity

remains unclear. Hopefully, future collections from both North America and

Europe will shed light on this subject.

SPECIMENS EXAMINED (for revised specimens see L. dicranellae)—ITALY. LOMBARDY,

Calvagese, Brescia, 300 m a.s.l., with Dicranella, Ditrichum, Riccia, and Anthoceros,

09.VI.2001 GM (MCVE 15993 as L. ascoboloides).

704 ... Eckstein & al.

Lamprospora cailletii Benkert, Z. Mykol. 53: 211. 1987.

Based on a collection from the French Alps, L. cailletii is characterized by

1-2 mm diameter yellow-orange apothecia with small fimbriate margins,

globose ascospores (14)15-16.5(17) um, ornamented by curved ridges

0.8-1.5 um wide and high forming a reticulum and additional warts, and

Tortella tortuosa as host (illustrations in Kristiansen 2006, Stdckli 2016).

The additional warts are sometimes difficult to see, and then the ascospore

ornamentation looks very similar to that of L. benkertii. It further differs from

L. benkertii by the small apothecial margins, slightly larger ascospores, lack of

secondary reticulum, and identity of the host.

Formerly L. cailletii was only rarely reported from France and Germany

(Benkert 1987), Norway (Kristiansen 2006), and Switzerland (Stockli 2016).

Newly, we report it from Slovakia. We doubt that the report on Ditrichum

flexicaule (Schwagr.) Hampe by Schumacher (1993) from France is actually

from this species.

SELECTED SPECIMENS EXAMINED—SLOVAKIA. ZILINA, Terchova, 1870 m SSW of St.

Cyril and Metod‘s church, Mala Fatra NP, 49.2417°N 19.0261°E, 935 m a.s.L, on soil

with Tortella tortuosa, 29.X.2016, ZE & MS (PRM 951726). SWITZERLAND. Jura,

Montfaucon, 47.2755°N 7.0793°E, 930 m a.s.l., host Tortella tortuosa, 18.X.2015 ES (B

70 0100014).

Lamprospora dicranellae Benkert, Z. Mykol. 53: 217. 1987.

Based on collections from the Czech Republic, Germany, Denmark, France,

Norway, and Sweden, L. dicranellae is characterized by orange apothecia 1-3

mm in diameter with conspicuous fimbriate margins, globose ascospores

(14)15-16.5(17) um, ornamented by curved ridges 1-2 um wide and high

forming a reticulum and additional warts (Fic. 7A). The host is not Dicranella

as assumed in the protologue but Ditrichum heteromallum and D. pusillum

(Eckstein & al. 2014, Vega & al. 2017). L. dicranellae differs from L. benkertii by

its larger apothecia with conspicuous fimbriate margins, the existence of many

prominent warts in addition to curved ridges on the ascospore surface, and

identity of the host.

Lamprospora dicranellae is a relatively widespread species with further

occurrences not mentioned in the protologue, e.g., in Austria and on the

Canary Islands and Madeira (Korf & Zhuang 1991) and in Great Britain (Yao &

Spooner 1995), the Netherlands (Brouwer 1999), Spain (Rubio & al. 2002), and

the USA (Wang & Kimbrough 1992).

Lamprospora benkertii sp. nov. (Europe) ... 705

SELECTED SPECIMENS EXAMINED—AUSTRIA. CARINTHIA, Nockberge, above

Zechneralm, 46.9494°N 13.7576°E, 1970 m a.s.L., 10. VIII.2015 JE42349 (B 70 0100988).

SALZBURG, Uttendorf, Staubachtal, on sandy soil near mosses, 18.VHI.2005 GM (K[M]

138534 as L. ascoboloides). GERMANY. THURINGIA, Eisenach, S Wartburg, 50.9581°N

10.2998°E, 280 m a.s.l., host Ditrichum sp., 27.1X.2017 JE51668 (B 70 0100989).

NORWAY. OppLAND, Vang, Tyinholmen, 1100 m a.s.l., host Ditrichum cf. pusillum,

23.VIII.1986 RK (TRH F-9757 as L. ascoboloides).

Lamprospora funigera McLennan & Cookson, Proc. Roy. Soc. Victoria 38: 70. 1926.

Based on two collections from the state Victoria in Australia, L. funigera is

characterized by apothecia 1-3 mm in diameter with well-developed fimbriate

margins, globose ascospores 12-13 um in diameter, ornamented by curved

ridges forming a reticulum. The host is unknown. It differs from L. benkertii in

larger apothecia with a conspicuous fimbriate margin and smaller ascospores.

So far L. funigera is only known from the type specimen (McLennan & Cookson

1926).

Lamprospora hispanica Benkert, Z. Mykol. 53: 227. 1987. Fics 6, 7B

This species is based on one collection from Spain. Studying the holotype

in MA we found some differences between the original description and our

observations later confirmed by many identical collections from different

countries. Because of the partly incorrect original description, we provide here

a more complete description based on our own observations (Fic. 6, 7B):

APOTHECIA gregarious, occasionally packed, on soil between shoots of

Aloina, 1-2(3) mm in diameter, at first spherical, becoming cup-shaped, finally

thick and discoid with a flat hymenium, sessile, with hyaline and flexuous

anchoring hyphae; MARGIN conspicuous, fimbriate to shaggy; HYMENIUM

varying from pale orange to orange, margin and outer surface paler.

Asci 250-450 x 20-28 um, cylindrical, 8-spored, unitunicate, operculate,

inamyloid, bifurcate at the base, arising from croziers.

Ascospores subglobose, hyaline, uniseriate, 16-19(22) x 15-18(20)

um (ornamentation included), Q: 1.03-1.06(1.1), always with a lipid drop

(10)11-13 um in diameter; ORNAMENTATION consisting of an alveolate

reticulum of the seaveri-type, with 5-8(10) meshes across ascospore diameter,

with the net’s density varying considerably between collections, meshes fairly

variable, often with lower and thinner ridges that may form a secondary net,

ascospore surface slightly roughened with occasionally minute stains, main

706 ... Eckstein & al.

ridges approximately 0.8-1.0 um wide and 0.6-0.8(1) um high, sometimes

with loops, anastomosing ridges not or rarely tapering at interjections.

PARAPHYSES filiform, straight, pluriseptate, branched at the base, gradually

enlarging towards the apex, terminal cell 50-120 x 4-8 um, distinctive

carotenoid pigment turning cyan to olivaceous in Lugol's solution, at times in

lipid bodies, numerous vacuolar bodies <4 um in size.

MEDULLARY EXCIPULUM textura intricata; ECTAL EXCIPULUM textura

prismatica-angularis; MARGO textura prismatica-porrecta, cells 36-120 x

10-30 um.

INFECTION on strong rhizoids of Aloina aloides (Koch ex Schultz) Kindb.

or Aloina ambigua (Bruch & Schimp.) Limpr. consisting of appressoria,

infection peg and haustoria; APPRESSORIA Clearly differentiated, attached to

the rhizoid becoming completely covered which anchoring hyphae, 30-60(75)

x 25-35(50) um in side view, mostly 2-septate, often multiple infection pegs

originating from one appressorium.

ECOLOGY & DISTRIBUTION—L. hispanica occurs on soil in open and

mostly warm habitats such as in dry grassland, in cemeteries, or along paths.

It does not seem to be rare in Spain (Ortega & Vizoso 1991, Martinez-Gil &

Caballero 2015). Additional collections have been made in Croatia, Cyprus,

France, Germany, Hungary, Portugal, and The Netherlands, with distribution

seemingly centered in the Mediterranean region with scattered occurrences

further north in the temperate zone. Given its known distribution, it is possible

that the species might occur in all Central and Southern European countries.

Lamprospora hispanica differs from L. benkertii by its larger apothecia with

conspicuous fimbriate margins, its slightly subglobose ascospores with more

uniformly wide ridges forming a denser net, its much larger appressoria, and

the identity of its host.

SELECTED SPECIMENS EXAMINED—CROATIA. Karuovac, Modrus rest area,

45.1301°N 15.2483°E, 500 ma.s.l., on soil with Aloina sp., 20.X1.2016 ZE (PRM 952410).

CYPRUS. Papuos, Peiya, 34.8936°N 32.3794°E, 440 m a.s.l, on soil layer on stone

wall, host Aloina sp., 28.11.2019 MV190228-07 (B 70 0100981). FRANCE. GIRONDE,

Cenon, Parc Palmer, 44.8611°N 0.5261°W, 55 m a.s.L., slope beside a path, host Aloina

ambigua, 06.1V.2018 MV180406-01 (B 70 0100978). HAUTE GARONNE, Toulouse,

cemetery of Terre-Cabade, 43.6067°N 1.4603°E, 170 m a.s.l., on soil between graves,

host Aloina ambigua, 14.11.2017 MV170214-03 (B 70 0100980). GERMANY. HEsseE,

Werra valley, Weinberg W Jestadt, 51.2188°N 9.9955°E, 180 m a.s.l., on soil in dry

grassland, host Aloina ambigua, 18.1.2014 JE34631 (B 70 0100990); LowER Saxony,

N Tettenborn, 51.5716°N 10.5534°E, 320 m a.s.l., in an old quarry on loamy soil with

707

Lamprospora benkertii sp. nov. (Europe) ...

H. B 70 0100980). A, B. Apothecia on

soil with shoots of Aloina sp. C. Ascospores in water. D. Ascospores stained with cotton blue.

Fic. 6. Lamprospora hispanica (A. B 70 0100990;

E. Paraphyses and asci in water. F. Cross section of an apothecium. G. Medullary excipulum

next to the hymenium. H. Cross section of upper margin of an apothecium. Scale bars:

708 ... Eckstein & al.

gypsum, host Aloina aloides, 22.X1.2016 JE49371 (B 70 0100992). HUNGARY. FEJER,

Vertes Mts., Csakvor, $z010-K6. 47.3804°N 18.4374°E, 200 m a.s.l., host Aloina ambigua,

05.11.2016 CN7751 (private herbarium CN). PORTUGAL. LisBoa, Cémiterio

do Alto de Sao Joao, 38.7303°N 9.1206°W, 70 m a.s.l., on soil between graves, host

Aloina ambigua, 25.1.2014 MV140125-13 (B 70 0100995). SPAIN. ARAGON, Teruel,

Calle Bajo Los Arcos, 40.3458°N 1.1058°W, on soil in front of a house, host Aloina

ambigua, 30.IX.2015 RTL15093001 (B 70 0100985); La Rioja, Logrono, Parque

La Grajera, 42.4464°N 2.5078°W, 440 m a.s.l., on soil between trees of Populus sp.,

05.XII.2014 RMG (B 70 0100979); MapRID, Aranjuez, 06.11.1975 MdIT (MA-Fungi

2359, holotype); MALLORCA, Inca, Parque Llubi Torradas, 39.7064°N 2.9967°E, 80 m

a.s.l., below a tree, host Aloina sp., 01-XII.2017 MV171201-06 (B 70 0100983); Palma

de Mallorca, Placa del Rentadors, 39.5694°N 2.6389°E, 10 m a.s.l., amenity land, host

Aloina ambigua, 22.1.2017 MV170122-03 (B 70 0100986); Pais VAsco, Parque Ribera,

host Aloina ambigua, 23.1.2016 MV160123-02 (B 70 0100008); Urkiola, Calle Barrio

de Urkiola, 43.1003°N 2.6447°W, 730 m a.s.l., bank alongside the road, host Aloina

aloides, 19.V1.2018 MV180619-01 (B 70 0100998). THE NETHERLANDS. ZEELAND,

Oosterschelde-dam, south of Burgh-Haamstede, on Neeltje-Jans near Expo, 1 m a.s.l.,

on sandy recreation area, partly covered with gravel, host Aloina ambigua, 31.XII.2008

EB270 (private herbarium EB).

Lamprospora irregulariata K.B. Khare & V.P. Tewari,

Mycologia 70: 885. 1978.

Based on a collection from Mussoorie in the state Uttarakhand in northern

India, L. irregulariata is characterized by apothecia 2-3 mm in diameter,

a yellow-white hymenium, globose ascospores 13-16 um, ornamented by

thick curved ridges forming an incomplete reticulum. The host is not known.

The yellow white color of the hymenium set this species clearly apart from

L. benkertii and even raises doubts in its afhliation to the genus Lamprospora.

It is known only from the type (Khare & Tewari 1978).

Lamprospora leptodictya Dissing, Mycologia 73: 263. 1981.

Based on a collection from Greenland, L. leptodictya is characterized by

apothecia 0.8-1.8 mm in diameter with conspicuous fimbriate margins, an

orange hymenium, subglobose ascospores, 15-17 x 13-15.5 um, Q <1.1,

ornamented with curved, often blunt ending and tapering and only partly

anastomosing ridges 0.5-0.9 um wide and 0.5 um high (Fic. 7C), and its host

Aongstroemia longipes (Sommerf.) Bruch & Schimp. It differs from L. benkertii

by its subglobose ascospores, smaller ridges not forming a reticulum, and its

host.

Dissing notes A. longipes and Bryum sp. as accompanying bryophytes.

Benkert (1987) studied a later collection from the type locality and suspected

Lamprospora benkertii sp. nov. (Europe) ... 709

Fic. 7. SEM of ascospores of species similar to Lamprospora benkertii. A. L. dicranellae

(B 70 0100989); B. L. hispanica (B 70 0100998); C. L. leptodictya (B 70 0100009); D. L. norvegica

(B 70 0100021); E. L. paechnatzii (private herbarium TR 22VI2014, s.n.); F. L. seaveri

(B 70 0010018, holotype). Scale bars = 10 um.

Bryum sp. as host but could not actually find the infection. In specimens

from the French and Swiss Alps we confirmed A. longipes as the host. The

ornamentation is similar to that of L. paechnatzii with often short and blunt

ending ridges not or hardly forming a reticulate pattern.

Based on the few reports and the distribution of the host moss, L. leptodictya

is an arctic-alpine species, known from Greenland (Dissing 1981), Norway

(Schumacher 1993), France (Van Vooren 2017), and Switzerland.

710... Eckstein & al.

SPECIMEN EXAMINED—FRANCE. Savoy, Bonneval-sur-Arc, Ruisseau de

Céma,45.4250°N 7.0419°E, 2740 m a.s.l., host Aongstroemia longipes, 26.VIII.2015

MV20150826-01 (B 70 0100009). SWITZERLAND. BERN, Guttannen, Oberaarstausee,

46.5474°N 8.2769°E, 2306 m a.s.l., host Aongstroemia longipes, 18.VII.2014 ES (ZT Myc

61079).

Lamprospora norvegica Benkert, Aas & R. Kristiansen, Z. Mykol. 57: 196. 1991.

Based on collections from mainland Norway, L. norvegica is characterized

by apothecia 0.7-1.0(1.5) mm in diameter with conspicuous fimbriate margins,

an orange hymenium, globose to slightly subglobose ascospores, 13-14 x

12-13 um (Q <1.08), ornamented with curved ridges 0.3-0.7 um wide and

0.5 um high forming a reticulum (Fic. 7D). It differs from L. benkertii by

slightly subglobose ascospores, smaller ridges, and host.

Benkert & al. (1991) suspected Pohlia filum as host but could not actually

prove this. We studied collections from Norway and Switzerland and in two

collections, we could identify Ditrichum pusillum (Hedw.) Hampe as host. The

infection causes the swelling of infected rhizoid cells forming spherical galls.

Lamprospora norvegica is known from Norway (Benkert & al. 1991),

Svalbard (Schumacher 1993), and Switzerland.

SPECIMENS EXAMINED—NORWAY. BuSKEROD, Hol, Raggsteindalsvegen, 60.6666°N

7.8147°E, 1000 m a.s.l., host Ditrichum pusillum, 01.VII.2015 RK (TRH F-17572).

HORDALAND, Ulvik, Finse, near Finse railway station, 60.60°N 7.50°E, 1300 ma.s.L, host

probably Ditrichum pusillum, 24.V11I.2008 RK29/06 (TRH F-17573). SWITZERLAND.

BERN, Gadmen, Steingletscher, 46.7233°N 8.4340°E, 1930 m a.s.l., host Ditrichum

pusillum, 17.V1I.2014 ES (B 70 0100021).

Lamprospora paechnatzii Benkert, Z. Mykol. 53: 237. 1987.

Based on a collection from the German lowlands L. paechnatzii is

characterized by apothecia 1-2 mm in diameter, with low but conspicuous

fimbriate margins, a yellow to yellowish orange hymenium, globose ascospores

(14-16 um in diameter) ornamented with curved often blunt ending and

tapering ridges 0.3-0.8 um wide and high, short and often S-shaped, rarely

anastomosing and not forming a reticulum (Fic. 7E), and the host Bryum

archangelicum Bruch & Schimp. It differs from L. benkertii by smaller ridges

not forming a reticulum and the host.

Benkert reports Bryum sp. as the host; but our examination of the type

specimen revealed B. archangelicum as the host moss.

Lamprospora paechnatzii is known from Germany and the Czech Republic

(Benkert (1987) and Spain (Rubio & al. 2002).

Lamprospora benkertii sp. nov. (Europe) ... 711

SPECIMENS EXAMINED—GERMANY. BRANDENBURG, Bernau, Kiesgrube

Schwanebeck-Alpenberge, host Bryum archangelicum, V1I.1978 EP (B 70 0010013,

holotype); MECKLENBURG-WESTERN POMERANIA, Liitzow, gravel pit N Pokrent,

host Bryum sp., 22.V1.2014 TR (private herbarium TR); host Bryum sp., 05.VIIL2014

MV 140805-04 (B 70 0100018).

Lamprospora seaveri Benkert, Z. Mykol. 53: 241. 1987.

Based on many collections from Central and Northern Europe, L. seaveri

is characterized by orange apothecia 1-4 mm in diameter with conspicuous

fimbriate margins, an orange hymenium, globose ascospores 14-16 um in

diameter with an ornamentation consisting of fine ridges 0.3-0.8 um wide and

high forming a reticulum, often with a secondary reticulum within the meshes

(Fic. 7F), and the host Ceratodon purpureus. It differs from L. benkertii by its

larger apothecia with a conspicuous margin, smaller ridges, and host identity.

Benkert (1987) mentions the misapplication of the name Lamprospora

laetirubra (Cooke) Lagarde for this species; the type of the basionym Peziza

laetirubra Cooke is conspecific with Pulvinula cinnabarina (Fuckel) Boud.

Although the designated holotype of L. seaveri has Ceratodon purpureus as host,

Benkert (1987) also listed many collections on Bryum sp. among the paratypes.

As stated in the introduction, multiple unrelated hosts of a species often

indicate unresolved taxa. The ascospores of collections from C. purpureus and

from Bryum dichotomum are indeed very similar, and variations in ascospore

size and ornamentation partly overlap. Nevertheless, our suspicion of hidden

diversity is also supported by molecular data (Fic. 5, L. aff. seaveri), which

indicate an independent taxon on Bryum dichotomum. Because of mostly scant

collections on this host, description of L. aff. seaveri must be addressed in a later

study. For the meantime, we recommend that only collections on C. purpureus

be identified as L. seaveri.

SELECTED SPECIMENS EXAMINED—AUSTRIA. VIENNA, Central Cemetery, host

Ceratodon purpureus, 01.XI1.2014 MV141201-02 (B 70 0010019). GERMANY.

BRANDENBURG, Potsdam, slope along railway embankment at Berlin ring road

near bridge across Caputher Chaussee, host Ceratodon purpureus, 23.X11.986 DB

(B 70 0010018, holotype). SaxoNY-ANHALT, Galgenberg, NE Elbingerode, host

Ceratodon purpureus, 10.X.2015 GE(B 700100997). SCHLESWIG-HOLSTEIN, Appen,

host Ceratodon purpureus, 08.X1.2015 MV151108-01 (B 70 0100010). THURINGIA,

S Wipperdorf, 51.4463°N 10.6425°E, 240 m a.s.l., host Bryum dichotomum,

13.X.2017. JE51710 (B 70 0100993). MONTENEGRO. ANDRIJEVICA

Mounicipa.ity, Stavna plateau, 42.6988°N 19.6745°E, 1820 m a.s.l., between

mosses on the edge of the pathway on mountain pasture, host Ceratodon purpureus,

18.VI.2017 LJ (PRC 4581).

712 ... Eckstein & al.

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Lamprospora benkertii sp. nov. (Europe) ... 713

The main features of the mentioned species are summarized in TaBLE 2. To

facilitate identification, we provide a new key.

Key to species of Lamprospora with

seaveri-type (or similar) ascospore ornamentation

1. Hymenium yellow-white, apothecia 2-3 mm in diameter, little known

species from India (possibly not belonging to Lamprospora) ... L. irregulariata

I. Hymenium shades ‘of orange, 6). \di.24 ce ie ace tte Sa bike ee done oe ite o-edine els 2

2. Ascospores with main ridges 1-2 um wide and many prominent warts

between the ridges, host Ditrichum spp.................0 eee L. dicranellae

2. Ascospores with different ornamentation or host .............. 0. cece eee ee eee 3

3. Ascospores ornamented by often blunt ending ridges not or hardly

FORMING, ATSC CUTIES siace-e oi lacot dihince PAiine SBihne oAige ee hile sce 4 Nore Ae malleus 4

3. Ascospores ornamented by anastomosing ridges forming a reticulum ........... 6

4, Ridges of ascospore ornamentation strong, 1-2 um wide, ascospores globose,

13-15(-17) um, host probably Dicranella heteromalla.......... L. ascoboloides

4, Ridges of ascospore ornamentation weaker, 0.3-0.9 um wide, a

scospores globose or slightly subglobose .......... 0.0... cece eee eee eee 5

5. Ascospores mostly subglobose, 15-17 x 13-15.5 um, host Aongstroemia longipes,

intarctic=alpine NADItAtS MS 424. “Saihng gyiheceeaene ye Aitie-ea Aes 8 Bie ee L. leptodictya

5. Ascospores globose, 14-16 um, host Bryum archangelicum,

in the-témiperdte Zone Sac 34 shed ade seats +s file sa hie oe L. paechnatzii

6. Apothecia with conspicuous fimbriate margins, ascospores subglobose,

16-19(-22) x 15-18(-20) um, host Aloina spp............. 0.000. L. hispanica

6. Other combination of characters ..... 1... Lecce eee eee eee Ze

7. Main ridges of ascospore ornamentation 1 um wide or wider .................. 8

7. Main ridges of ascospore ornamentation 0.3-0.8(-1) um wide ................. 9

8. Main ridges of ascospore ornamentation 0.8-1.5 um wide with additional

warts (difficult to observe), host Tortella tortuosd...........000000 L. cailletii

8. Main ridges of ascospore ornamentation about 1 um wide often with a

secondary reticulum within the meshes, host Trichostomum crispulum,

southern-temperate distribution..................0 cece ee eae L. benkertii

9. Apothecia <1 mm, ascospores often slightly subglobose, 13-14 x 12-13 um,

host Ditrichum pusillum, in arctic-alpine habitats ............... L. norvegica

9. Apothecia mostly >1 mm, host and habitat different .....................004.8 10

10. Ascospores globose, 12-13 um,

little known species from Australia, host unknown ................ L. funigera

10. Ascospores globose, 14-18 um,

host Ceratodon purpureus or Bryum Spp....... 00. 11

714 ... Eckstein & al.

11. Ascospores globose, 14-16 um,

hest-Ceraidd on pur pureus- 22 tes gs toh tape on Rhy ee hy ee el Ee a L. seaveri

11. Ascospores globose or slightly subglobose, 14-18 um,

HOSE VPS Ptah ats. ea es state e tee tata ee le ed eae a eee a eae te ee L. aff. seaveri

There are several unpublished collections on different hosts with a seaveri-type

ascospore ornamentation that cannot be ascertained as conspecific with any

of the above-mentioned species. Because of these scant collections, no further

details are provided here, but these unresolved specimens indicate a higher

diversity in this group than previously thought.

Both morphological and molecular data confirm the very high host

specificity of the bryophilous Pezizales. All evaluated species grow only on one

particular moss species or on very closely related species as is the case with

L. hispanica with hosts Aloina aloides and A. ambigua. In L. seaveri, which

was reported from multiple hosts, the phylogenetic analysis clearly shows the

presence of at least one other species on Bryum dichotomum (Fic. 5).

Although an identification of species can be undertaken by morphological

characters only (TABLE 2), differences between species are sometimes subtle

and variations of some features can overlap. The easiest way to differentiate

between species is to consider the ecology and host identity. Specimens from

hitherto unknown hosts, or with ascospores differing from those of known

species on a particular host, are likely candidates for new species.

Acknowledgements

We express our gratitude to Giinter Eckstein (Germany), Karen Hansen

(Sweden), Roy Kristiansen (Norway), Rubén Martinez-Gil (Spain), Csaba Németh

(Hungary), Torsten Richter (Germany), Elisabeth St6ckli (Switzerland), Raul Tena

Lahoz (Spain) for sending samples and documentation, to Michal Sochor (Czech

Republic) for molecular analysis, and to Markus Wagner for the improvement of the

English language. We also thank the curators of B, K, MA, MCVE, NY, TRH for the

opportunity to study specimens, and the staff of the Johann-Friedrich-Blumenbach-

Institute for Zoology and Anthropology in Gottingen for allowing us to take SEM

photographs using their equipment. The expert review comments and suggestions

by Peter Dobbeler (Ludwig-Maximilians-Universitat, Miinchen, Germany) and

Nicolas Van Vooren (Lyon, France) are greatly appreciated.

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MY COTAXON

October-December 2021—Volume 136, pp. 719-723

https://doi.org/10.5248/136.719

Pertusaria wui sp. nov. on bamboo from Yunnan, China

JIARONG ZHANG’, LIN LIU’, XIANDONG XUE’, QIANG REN?

' College of Life Sciences, Shandong Normal University,

No. 88 East Wenhua Road, Jinan 250014, China

’ State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences,

No. 1 West Beichen Road, Beijing 100101, China

“CORRESPONDENCE TO: rendaqiang@hotmail.com

ABSTRACT—A new species Pertusaria wui is described from bamboo in Yunnan Province,

China. Diagnostic characters include a thin, shiny, off-white, continuous and crustose

thallus, numerous (mostly conical) verrucae with punctiform ostioles, uniseriate ascospores

of 47.5-62.5 x 25-30 um, 8-spored asci, and major lichen compounds lichexanthone and

stictic acid.

Key worps—lichenized Ascomycota, Pertusariaceae, Pertusariales, taxonomy

Introduction

After molecular and allied data by Lumbsch & Schmitt (2001) indicated that

Pertusaria DC. s.lat. was polyphyletic, Schmitt & Lumbsch (2004) identified

three well-supported clades: /Pertusaria s.str., /Varicellaria, and /Variolaria.

Archer & Elix (2016) divided Pertusaria DC. s.lat. into three subgenera, P. subg.

Monomurata A.W. Archer, P. subg. Pionospora Th. Fr., and P. subg. Pertusaria

based on morphological and chemical characters. However, the molecularly

supported clades were elevated to the genera as Pertusaria DC. s.str., Varicellaria

Nyl., and Lepra Scop. (Schmitt & al. 2012; Hafellner & Turk 2016; Wei & al.

2017), genera currently accepted by lichenologists (Archer & Elix 2018; Zhou

& Ren 2018; Ren 2019). Pertusaria s.lat. in China was first annotated by Zhao

& al. (2004), additional new taxa and new records have been published (Ren

& al. 2008, 2009a,b; Ren & Kou 2013; Ren 2013, 2014, 2015; Ren & Zhao 2014;

Zhang & Ren 2016). As part of a continuing study of Pertusaria s.str. from

720 ... Zhang & al.

China, we re-examined previously unidentified specimens to discover that a

single KUN specimen represented a new species, described here as Pertusaria

wui.

Materials & methods

The examined specimen is deposited in the Herbarium, Kunming Institute of

Botany, Chinese Academy of Sciences, Kunming, China (KUN). It was examined

microscopically under an Olympus SZX16 dissecting microscope and a Zeiss

Axioskop 2+ compound microscope; chemical substances were identified by thin-

layer chromatography using solvents A, B, and C (Elix 2014) and compared with

authentic samples. We photographed the specimen with a Nikon D700 fitted with a

Micro-Nikkor 105 mm f/2.8G AF-S VR IF-ED lens.

Taxonomy

Pertusaria wui Q. Ren, sp. nov. Fig. 1

EN 570761

Differs from Pertusaria tetrathalamia by its 8-spored asci and its lack of

4,5-dichlorolichexanthone.

TypE—China. Yunnan Province, Nanjian County, Mt. Dadian, 24.85°N 100.57°E, on

bamboo, 21 Mar. 2012, L.S. Wang 12-32980, X.Y. Wang & D. Liu (Holotype, KUN).

ErymMoLocy—honoring the late Prof. Jinong Wu, who contributed greatly to Chinese

lichen research.

THALLUuS off-white, thin; surface smooth, shiny, continuous, lacking isidia

and soredia. APOTHECIA conspicuous, concolorous with thallus, numerous,

verruciform, frequently conical and smooth, generally dispersed and only

locally fused, 0.5-0.8(-1.5) mm in diameter. Ostioles conspicuous, black,

punctiform, 1-3(-5) per verruca, surrounded by a white border, mostly level

with verruca surface but occasionally papillate. AscospoREs 8 per ascus,

longitudinally uniseriate, hyaline, ellipsoid, 47.5-62.5 x 25-30 um; ascospore

wall smooth, c. 5 um thick except for thicker (c. 10 um) end wall. Pycnrp1 not

observed.

CHEMISTRY—Cortex UV+ yellow-orange, all chemical tests negative.

Medulla K+ yellow, C-, KC-, Pd+ yellow, then orange. Lichexanthone and

stictic acid detected.

DisTRIBUTION—Currently known only from the type locality in Yunnan.

CoMMENTS—Pertusaria wui is characterized by the numerous conical

verrucae, asci with 8 uniseriate smooth-walled ascospores, and the presence

of lichexanthone and stictic acid. The new species closely resembles

P. tetrathalamia (Fée) Nyl. from North America and China (Dibben 1980;

Pertusaria wui sp. nov. (China) ... 721

‘a *. ; —

Fic. 1. Pertusaria wui (KUN - Wang 12-32980). A. Thallus with substratum; B. Apothecia, detail;

C. Asci with mature or immature ascospores, arrows pointing to eight uniseriate and immature

ascospores within one ascus; D. Mature ascospores. Scale bars: A, B = 1 mm; C, D = 30 um.

722 ... Zhang & al.

Zhao & al. 2004) in also presenting conspicuous black ostioles and mostly

simple verrucae, but P. tetrathalamia has 4-spored asci and contains stictic

acid and 4,5-dichlorolichexanthone.

Beside the new species, only four other Pertusaria species possess 8-spored

asci and contain lichexanthone and stictic acid as major lichen substances.

Among them, Pertusaria nigrata Kremp. from Brazil (Krempelhuber 1876)

and Sri Lanka (Weerakoon & Aptroot 2014) is most similar to the new species

but differs in having longer ascospores (75-90 um). The other three species are

P. dehiscens Mill. Arg., P. fosseyae A.W. Archer & al., and P. parnassia Vain.

Their differences from P. wui are indicated in the key below.

Key to Pertusaria spp. having 8-spored asci and

lichexanthone + stictic acid as major lichen substances

1. Thallus sorediate (spores 2-seriate, 98-111 um long);

Kahuzi & Virunga mountain ranges in East Africa's Albertine Rift ... P fosseyae

Mpa les GOES UALS Brn. aM ca dete a. las Recah ad ied at Ae Pa Se PS PRE An le sect ee Z

2. Spore walls rough (spores 110-180 um long);

Gira del ibe: WESTIE cites, ag dicta, 3 ise eg Lublin 4g Std ber a Aidt eee 3 Athan atts P. parnassia

2 SPOKE WALIS SURO O Uae fees) cpoteeaet aco nttel ohio tees ore iat cided esas deeb ayacyapolancse ted mesa de mgonds eens 3

3. Spores 2-seriate (90-140 um long);

Australig, Brazil, chile: MGs Ti aMkad, sxe chs ceva ets save ahs serarahs Boe P. dehiscens

Be SPOTES UH SOR ALS ihe ues it wea bs Nae Mecca Widget Be Hagel Fae lag Wadden Yaa Wat Wg 4

4. Spores 75-90 um long (Brazil & Sri Lanka) .......... 0... eee eee eee P. nigrata

Ar Spores: 47 5=G2rouim Ome CCHIT is 6tats BiG at MS aes yea 5 UBS ae P wui

Acknowledgments

We appreciate the help of the curator (Prof. Zhuliang Yang) who loaned

material for this study. The authors thank Dr. Man-Rong Huang (Beijing Museum

of Natural History, PR. China) and Dr. Huajie Liu (Hebei University, China) for

pre-submission manuscript review. Special thanks go to Mr. Lisong Wang (KUN).

This study was financially supported by the National Natural Science Foundation of

China (32070011).

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Pertusaria wui sp. nov. (China) ... 723

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China. Bryologist 112: 394-396. https://doi.org/10.1639/0007-2745-112.2.394

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chemistry as an important systematic character set in lichen-forming ascomycetes. Molecular

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Schmitt I, Otte J, Parnmen S, Sadowska-Des AD, Liicking R, Lumbsch HT. 2012. A new

circumscription of the genus Varicellaria (Pertusariales, Ascomycota). Mycokeys 4: 23-36.

https://doi.org/10.3897/mycokeys.4.3545

Weerakoon G, Aptroot A. 2014. Over 200 new lichen records from Sri Lanka, with three new

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133: 71-77. https://doi.org/10.5248/133.71

MY COTAXON

October-December 2021— Volume 136, pp. 725-737

https://doi.org/10.5248/136.725

Clavaria cystidiata sp. nov. from India

K. KRISHNAPRIYA & T.K. ARUN KUMAR

Department of Botany, The Zamorins Guruvayurappan College,

Kerala 673 014, India

*CORRESPONDENCE TO: tkakumar@gmail.com

ABSTRACT—A new species, Clavaria cystidiata, was collected and described from Kerala

State, India. The species is proposed as new based on morphological and molecular evidence.

Clavaria cystidiata differs from other closely related Clavaria species by its garlic smell,

yellowish basidiomata, clamp connections at the base of basidia, and by the prominent

presence of versiform cystidia. Sequence analyses of the nuclear rDNA internal transcribed

spacer (ITS) region supports phylogenetic separation of C. cystidiata within Clavariaceae.

Keyworps—Agaricomycetes, alliaceous, biodiversity, Agaricales, coral fungi

Introduction

Clavaria Vaill. is a basidiomycetous genus, comprising about 178 described

species (Roskov & al. 2020). The genus has a cosmopolitan distribution.

Representatives grow primarily as saprobes and (exceptionally) as associates of

soil algae and ericoid plants (Corner 1950; Englander & Hull 1980, Olariaga &

al. 2015). Basidiomata are simple (rarely branched), cylindrical to club shaped,

and whitish, yellow, orange, pink, violet, or brown to black in color. Holobasidia

bear hyaline spores. Characteristics of the basidia, hyphae, and presence and

location of clamp connections have been used in infrageneric classifications.

Basidiomatal color has been used both as a key taxonomic character for

delimiting species and informally grouping species within the genus (Corner

1950, 1970; Knudsen 1997; Petersen 1978, 1988; Roberts 2007; Kautmanova &

al. 2012). Kautmanova & al. (2012) analyzed the phylogeny of dark Clavaria

species (with grey, brown, and black basidiomata) using LSU nrDNA sequence

data. Olariaga & al. (2015) studied yellow Clavaria species with clamped basidia

726 ... Krishnapriya & Kumar

and inferred species relationships based on ITS and LSU nrDNA sequence

analyses.

Similarly, the distinctive smell of basidiomata has served as a distinguishing

taxonomic character in the genus. Basidiomata of some Clavaria species are

known to emit a distinctive alliaceous smell, and Petersen (1988) used garlic

smell as one identifying character in his keys to the New Zealand clavarioid

species. Clavaria falcata Pers., C. foetida G.F. Atk., C. fuscata Oudem., and

C. redoleoalii R.H. Petersen are the four known garlic-smelling species of

Clavaria.

During an exploratory survey of the clavarioid fungi of Kerala, India, we

encountered basidiomata of a garlic-smelling yellowish clavarioid fungus.

Detailed examination revealed this collection as unique from all other

described Clavaria species. Details of our morphological and molecular

examination accompany the proposal of a new species. A key to Clavaria

species reported from India is also provided.

Materials & methods

Morphological characterization

Fruit bodies were collected from Kerala State, India, during August 2017. Fresh

material was examined macroscopically and tested with FeCl, and KOH. Dried

materials were examined microscopically and stained using aqueous solutions of 3%

phloxine and 1% Congo red mounted in 5% aqueous KOH. Twenty basidiospores

were measured to calculate dimensions, mean, and standard deviation for length

and width, range of spore quotient (Q, length/width ratio) and its mean value (Q_).

Basidiospore reactivity to Melzer’s reagent and cotton blue was noted. The holotype

was deposited at the Central National Herbarium, Kolkata, West Bengal, India (CAL),

and an isotype was deposited at the Zamorin’s Guruvayurappan College Herbarium,

Kozhikode, Kerala, India (ZGC).

Molecular characterization & phylogenetic analysis

Genomic DNA was extracted from dried basidiomata, using the procedure of

Izumitsu & al. (2012). The ITS region was amplified by PCR using primers ITSIF

and ITS4 (Gardes & Bruns 1993, White & al. 1990). PCR product was purified

using ExoSAP-IT treatment and sequenced using BigDye Terminator v3.1. The

newly generated sequence was deposited in GenBank (www.ncbi.nlm.nih.gov) with

accession number MK751792.

The newly generated sequence was subjected to a BLAST search in the GenBank

nucleotide database for finding taxa with close sequence similarity. A data matrix was

constructed with the newly generated sequence and ITS sequences retrieved from

GenBank of 47 representative sequences in Clavariaceae (TABLE 1). All Clavaria

species with ITS sequences available in GenBank were included in the dataset.

Clavaria cystidiata sp. nov. (India) ... 727

TABLE 1. Taxon and GenBank accession numbers for the ITS sequences used in

phylogenetic analyses.

TAXON

Clavaria acuta

Clavaria amoenoides

Clavaria argillacea

Clavaria asterospora

Clavaria atrofusca

Clavaria calabrica

Clavaria californica

Clavaria citrinorubra

Clavaria crosslandii

Clavaria cystidiata

Clavaria falcata

Clavaria flavipes

Clavaria flavostellifera

Clavaria fumosa

Clavaria fuscata

Clavaria greletii

Clavaria greletoides

Clavaria griseobrunnea

Clavaria incarnata

Clavaria redoleoalii

Clavaria rosea

Clavaria rubicundula

Clavaria sphagnicola

Clavaria tenuipes

Clavaria tyrrhenica

Clavaria zollingeri

Clavicorona taxophila

Clavulinopsis amoena

Clavulinopsis cf. helvola

Clavulinopsis fusiformis

Clavulinopsis laeticolor

Clavulinopsis miyabeana

Clavulinopsis sulcata

Camarophyllopsis atrovelutina

Camarophyllopsis phaeophylla

Camarophyllopsis rugulosa

Camarophyllopsis atropuncta

Camarophyllopsis schulzeri

Hyphodontiella multiseptata

Mucronella sp.

Mucronella bresadolae

Ramariopsis flavescens

Ramariopsis kunzei

Ramariopsis crocea

Ramariopsis pulchella

Scytinopogon havencampii

GENBANK NO.

AY228353

MF972891

KC759438

KC759440

HQ606080

MF972889

HQ179660

HQ179661

KC75944

MK751792

KC759445

KC759451

KC759462

MK427065

KP257128

MF503244

MF503243

NR158336

KC759452

MF664111

MK909560

MK578690

KC759456

KC759457

MF972890

MH016820

AF033344

MK427063

KT275650

KM248914

EU118618

MK427059

MK427060

KU882900

MK139805

NR_119896

HQ662165

GU187556

EU118634

HQ533013

MH409972

DQ384591.1

NR_119913

MK616542

MK607557

KX812470

NR151488

STRAIN/HERBARIUM NO.

F14294

AMBO018217

K(M)126733

BIO-Fungi 12390

BRACR 13264

ZT Myc 58697

TENN:026785

TENN:040464

BIO-Fungi 12762

CAL 1769

AB0532

OJ362006

BRACR16695

Zp-2225

TENN065665

ERRO 2014102101

ERRO 2009070701

12566

BIO-Fungi 12560

PDD105311

TUR 201239

6603126

BRNM 747282

K(M) 146565

ZT Myc 58698

FLAS-F-60642

71850

ZP-2400

SE-2015

2718

EL 8/00

ZP.2118

ZP-2119

TL2014-682591

1ERRO 2013112901

TENN 023664

4G4-2010

GG091005

Ryberg 021022 (GB)

PDD95742

strain 1214

F15204

TENN 027570

AMB n. 17485

302989

MCCNNU00981

SFSU DED8300

LOCATION

Italy

England

Spain

Norway

Italy

USA

Australia

Spain

India

Wales

Austria

Spain

China

USA

Spain

New Zealand

Finland

USA

Czechia

England

Italy

USA

Finland

China

Denmark

France

USA

Ireland

Wales

Sweden

New Zealand

USA

Italy

USA

China

728 ... Krishnapriya & Kumar

Scytinopogon havencampii Desjardin & B.A. Perry (Hydnodontaceae, Trechisporales)

was chosen as the outgroup taxon following Birkebak & al. (2013). There were a total

of 717 positions in the final dataset. The aligned nrITS data matrix [ID: 26259] was

deposited in TreeBase. DNA sequences of the ITS dataset were aligned automatically

with MUSCLE in MEGA (Kumar & al. 2018) and then manually edited using the same

program.

Phylogenetic analyses were conducted using Maximum Likelihood (ML) and

Bayesian analysis (BA) methods. ML analysis was done with Tamura-Nei model

(Tamura & Nei 1993) in MEGA. Initial tree(s) for the heuristic search were obtained

automatically by applying Neighbor-Join and BioNJ algorithms to a matrix of

pair wise distances estimated using the Maximum Composite Likelihood (MCL)

approach, and then selecting the topology with superior log likelihood value. The tree

with the highest log likelihood value (—16331.38) was selected. The tree was drawn to

scale, with branch lengths measured in the number of substitutions per site. BA was

conducted using MrBayes 3.2 (Ronquist & al. 2012). Bayesian phylogenetic inference

was done with a T92+G model with discrete gamma distributed substitution rates

for the sequence dataset. The best-fit likelihood model of evolution was estimated in

MEGA. Multiple independent analyses were run from random starting trees for four

million generations, with trees saved every 100 generations, using four chains and a

burnin fraction of 0.25.

Taxonomy

Clavaria cystidiata Krishnapriya & T.K.A. Kumar, sp. nov. Figs 1, 2

MB 832680

Differs from other garlic-smelling, yellowish Clavaria spp. by its prominent versiform

cystidia and its bi-sterigmate basidia with basal clamp connections.

Type: India, Kerala State, Kozhikode District, Madapally Govt. College Campus,

9.4698°N 76.5902°E, altitude 11 m, on soil, solitary and in gregarious clusters among

dead and decaying leaf litter, 01 August 2017, K. Krishnapriya (Holotype, CAL 1769;

Isotype, ZGCKP59; GenBank MK751792).

EryMo_oey: cystidiata, referring to the prominent cystidia, the primary distinguishing

morphological character of the species.

BASIDIOMATA $110 mm long, <3 mm thick, simple, unbranched, cylindrical,

solid when young becoming hollow with age; terete in cross section; apex

acute, narrowing towards base; glabrous; off white to pale yellow when young,

becoming yellowish when mature, dark yellow to pale orange towards the apex;

context fleshy, with strong garlic odor; no positive reaction in FeCl,and KOH.

Basip1ospores 7-10 x 5-8 um (Q=1.1-1.5 um, Q =1.2 um), broadly

ellipsoid, contents granulate and guttulate (primarily one single large oil

droplet), smooth, thin-walled, hyaline, apiculus prominent (<1 um long),

inamyloid, cyanophilic in cotton blue. BAstp1a 50-60 x 7-10 um, aguttulate

Clavaria cystidiata sp. nov. (India) ... 729

a? . OO ere?

Fic. 1. Clavaria cystidiata (holotype, CAL 1769): Basidiomata. Scale bar=10 um.

to multiguttulate, cylindrical to clavate with basal clamp connection,

sterigmata 1-2(<5) um long. Cystrp1a abundant, 21-75 x 10-21 um,

versiform (predominantly cylindrical, clavate, lageniform, broadly clavate),

many with <27 um long apical protrusions, thin-walled, inamyloid, hyaline.

730 ... Krishnapriya & Kumar

Fic. 2. Clavaria cystidiata (holotype, CAL 1769). A. Basidiospores; B. Basidium with sterigmata;

C. Basidium with basal clamp connection; D-L. Cystidia. Scale bars: A, C= 8 um; B, D-L=10 um.

HyMENIUM 20-30 um wide. SUBHYMENIUM 50-60 um wide. CONTEXT

hyphae parallel, septate, inflated, 3-25 um wide, hyaline to pale yellow, thin-

to slightly thick-walled (<1 um), cyanophilic, inamyloid. Hyphal clamp

connections absent.

ECOLOGY & DISTRIBUTION: On soil. Both solitary and in gregarious

groups among leaf litter.

Clavaria cystidiata sp. nov. (India) ... 731

Molecular analysis

Sequencing of ITS region of the specimen of Clavaria cystidiata yielded

633 base pairs. The GenBank BLAST search of this newly generated sequence

produced sequence identities of only 83% with C. fuscata, 80% with C. falcata,

and 79% with C. redoleoalii, the closest BLAST hits. The ML (Fic 3) and BA

(Fic 4) phylogenetic trees were congruent with respect to the majority of clades

and taxa. Clavaria cystidiata clustered in a clade with the three garlic-smelling

taxa (C. fuscata, C. redoleoalii, C. falcata) and three non-garlic-smelling species

(C. greletoides, C. californica, C. tenuipes) with 85% ML bootstrap support (BS),

and 0.86 BA posterior probability (PP) support. The new species, C. cystidiata

resolved as sister to C. fuscata, receiving low (50% BS) value in the ML tree

but strong (PP 1) support in the BA tree. The yellow-coloured Clavaria species

included in the analyses—C. amoenoides Corner & al., C. argillacea Pers.,

C. citrinorubra R.H. Petersen, C. flavipes Pers., C. flavostellifera Olariaga & al.,

C. inaequalis (Pers.) Corner, C. sphagnicola Boud., C. redoleoalii—did not form

a monophyletic group. The monotypic Clavicorona taxophila (Thom) Doty

clustered with Clavaria crosslandii Cotton (84% BS, 0.94 PP). Camarophyllopsis

species included in the study were recovered in two separate clades in both

Bayesian and ML analyses. Hyphodontiella multiseptata A. Strid fell into a sister

clade to Ramariopsis and Clavulinopsis, forming a well-supported monophyletic

group, distinct from the Clavaria-Camarophyllopsis—Clavicorona clade in the

ML analysis. In the BA analysis, H. multiseptata fell into the Ramariopsis clade.

Mucronella forms a distinct clade from the rest (79% BS, 0.84 PP).

Discussion

Clavaria cystidiata is recognized by its yellowish colour, strong garlic smell,

basal clamp connection in basidia, presence of cystidia, large spores, and

inflated hyphae. The presence of strong garlic odor and prominent cystidia

are key characters which differentiate C. cystidiata from other yellow coloured

Clavaria species. Corner & al. (1956) proposed a yellow Clavaria species,

C. amoenoides from India. Its yellow colored basidiomata show similarity

with C. cystidiata macroscopically but the absence of clamp connection at

the base of basidia, absence of cystidia, and the absence of garlic odor (Thind

1961) makes it different.

Clavaria cystidiata closely resembles C. fuscata, C. foetida, C. redoleoalii, and

C. falcata morphologically (TABLE 2). During BLAST search in NCBI’s GenBank

nucleotide database C. cystidiata shows only 83% similarity with C. fuscata.

Morphologically C. fuscata differs from C. cystidiata in having white coloured

732... Krishnapriya & Kumar

87%

93%

92%

-— MK427065 Clavaria fumosa

83%

89% | ‘—— MF972689 Clavaria calabrica

82% '— MK578690 Clavaria rubicundula

a5% MHO16820 = Clavaria zollingeri

71%

MF972891 Clavaria amoenoides

97%

85% 8%

95%

91% 67%

89%

83%

81% |30%

84%

pons )K751792 Clavaria cystidiata®

50%

‘—— KP257128 Clavaria fuscata @

MF664111 Clavaria redoleoalii @

KC759445 Clavaria falcata @

adv | [————. MF 503243 Clavaria greletoides

88% | 73%

HQ179660 Clavaria califomica

KC759457 = Clavana tenuipes

74%

73% “—— NR 119896 Camarophyllopsis rugulosa

'—— HQ662165 Camarophyllopsis atropuncta

HQ606080 Clavaria atrofusca

72%

NR 158336 Clavaria griseobrunnea

r KC759441 Clavaria crossiandii

85%

'- AF033344 Clavicorona taxophila

HQ179661 Clavaria citrinorubra

65%

—— KC759448 Clavaria flavipes

90%

r-——— KC759439 Clavaria argillacea

90%

|____ KC759456 Clavaria sphagnicola

MK909560 Clavaria rosea

MF503244 Clavaria greletii

82%

87%

KC759440 Clavana asterospora

84%

KC759452 Clavaria incamata

MF972890 Clavaria tyrrhenica

78%

KC759462 Clavaria flavostellifera

£U118634 Hyphodontiella multiseptata

NR 119913 Ramariopsis flavescens

——— KX812470 Ramariopsis pulchella

MK607557 ~=Ramariopsis crocea

MK616542 Ramariopsis kunzei

KT275650 Clavulinopsis cf. helvola

49%

78% EU118618 Clavulinopsis laeticolor

— KM248914 Clavulinopsis fusiformis

79%

-—— ™MK427059 Clavulinopsis miyabeana

80% | -— MK427060 Clavulinopsis sulcata

73%

'——. MK427063 Clavulinopsis amoena

HQ533013 Mucronella species

DQ384591 Mucronella bresadolae

75%

‘_—— MH409972 Mucronella species

NR 154418 Scytinopogon havencampit

1DQ202267 Clavaria inaequalis

t——— _MK139805 Camarophyllopsis phaeophyila

AY228353 Clavaria acuta

GU187556 Camarophyllopsis schulzeri

KU882900 + Camarophyllopsis atrovelutina

Camarophyllopsis

| Clavicorona

Camarophyllopsis

Hyphodontiella

Ramaropsis

Clavulinopsis

Mucronella

Clavaria

733

Clavaria cystidiata sp. nov. (India) ...

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Fic. 3 (left). Maximum likelihood tree showing phylogenetic placement of Clavaria cystidiata

(shown in bold and with a thickened branch). ML BS values are shown. Sequences are identified

by GenBank accession number.

Fic. 4 (above). Bayesian analysis consensus tree for Clavaria cystidiata and related taxa, with PP

values shown on the branches. Four garlic-smelling species are indicated with asterisks. Sequences

are identified by GenBank accession number.

734 ... Krishnapriya & Kumar

TABLE 2. Morphological comparison between Clavaria cystidiata and

other garlic-smelling Clavaria spp.

C. cystidiata C. falcata C. foetida C. fuscata C. redoleoalii

BASIDIOME Pale yellow to White to Maize yellow White White

COLOR yellow cream

HyYPHAL DIAM 3-4um, — 6-20 3.5-12 4-12

(um) inflated <25

CYSTIDIA Present Absent Absent Absent Absent

SPORE SIZE 7-10 x 7-12 x 5.4-7.2 X 8-10 x 6.1-7.6 x

(tm) 5-8 4-9 3.2-4.0 4.5-6 5.4-6.5

BASIDIAL Present Absent Absent Absent Present

CLAMP

STERIGMATA 2 4 4 2 4

RANGE Tropical Tropical & Temperate Temperate Temperate

Temperate

basidiomata, absence of basal basidial clamp connections and cystidia, and very

small basidia (22-30 um, Corner 1950). According to Coker (1923), the spore

size of C. fuscata ranges from 4.4-6 x 8-10 um. The color of the fruit body and

strong garlic odor of C. foetida is similar to that of C. cystidiata. But spore size

(5.4-7.2 x 3.2-4.0 um), presence of four sterigmata, absence of cystidia, and

absence of basal clamp connections of basidia in C. foetida (Petersen 1988)

separates the two. Unfortunately, C. foetida was unavailable for DNA extraction

and sequencing for our molecular analyses, as the use of the only known and

available single basidiomata of the species deposited in a public herbarium

was restricted. Hence, this garlic-smelling taxon could not be included in this

study. C. redoleoalii is a garlic-smelling Clavaria, having abundant crystalline

material in tramal hyphae, with secondary septations and four sterigmata

(Petersen 1988). C. redoleoalii also lacks cystidia. These characters differentiate

C. cystidiata from C. redoleoalii. Sequence of C. redoleoalii showed only 79%

similarity with C. cystidiata during the BLAST search. Another garlic-smelling

taxon, C. falcata showed only 80% sequence similarity with C. cystidiata.

Morphologically, it differs from C. cystidiata in having white to cream colored

basidiomata, absence of basidial clamp connections, presence of four-spored

basidia, and absence of cystidia. C. cystidiata is unique from all the described

species of Clavaria by the presence of its prominent versiform cystidia.

Clavaria cystidiata sp. nov. (India) ... 735

Out of the eight Clavaria species described so far with yellowish

basidiomata, only one species (C. amoenoides) has previously been reported

from India (Verma & Pandro 2018). The phylogeny of yellow Clavaria species

has been studied before. Molecular investigations (Olariaga & al. 2015;

Birkebak & al. 2013, 2016) had proved that they do not form a monophyletic

group. Our ML and BA studies also support these observations. C. cystidiata

now ranks as the fifth garlic-smelling species of Clavaria to be collected and

reported globally. There are no reports of garlic-smelling Clavaria species

from India to date. Interestingly, our molecular analyses (ML and BA)

recovered all taxa with garlic-smelling basidiomata together as a separate

group. However, three non-garlic-smelling taxa also are settled in this

group. Phylogenetic placement of all other taxa of Clavariaceae, included

in our analyses, conforms to the results of previous studies (Birkebak & al.

2013, 2016; Olariaga & al. 2015). Our morphological studies and molecular

analyses reveal the distinction and phylogenetic placement of C. cystidiata

within the Clavariaceae. Based on these data, we propose the species as new

to science.

Key to the Clavaria species from India

Sasi TOM ate WPAN YC Ne aa ar ct tle ate ele ch Sd Ba Rd Sh ek ch ted SL 2

Ty Basidionvata Wn Deane eke len ke le We Ske bead a aie to Anh Nas eed 3

2. Basidiomata purple to pinkish purple; frequent to occasional branching C. zollingeri

2. Basidiomata white; always with crowded branches ................. C. jacquemontii

3. Loop-like clamp connections present at basidial bases ..................200005 +

3. Loop-like clamp connections absent from basidial bases....................04. 5

4vBasidiomata with garlic Odor. 7. 92h.'. Babies. Bagnirs. Page (os Daye fae eegs lot Pept C. cystidiata

4, Basidiomrata avout Garlie: OGM 2's saris tox arc ttavalste sarats travathe tava a wavartey ok tay 3 6

5; Basicioiiatany ClO w= 3 08.902 15.5% dnt 2s WMS 20d. 8RR) LotR! EBB ULB Be C. amoenoides

5. Basidiomata not yellow; brown, white, or grey ........ 0... cece cece eee eee ee

6. Basidiomata pink; basidiospores 6-7.5 x 4-4.5 um, ellipsoid;

hiphae notsecondarily septate ise i. ose ee vk ee yee cay ¥ wean ya C. incarnata

6. Basidiomata white; basidiospores 8-11 x 6-8 um, globose;

hyphae seconvatily septateus ss higes hose wh tage vee done Foe 2 HEM ote C. acuta

7. Basidiomata brown; basidiospores 5.5-7.5 x 2.5-3.2 um;

basidia 25-40 x 5.5-8 um; hyphae 2-13 um diam .................. C. indica

7 Basidionvata- WIVte On OTEV.., 2 dice e a dive 3d neg d ceed d weg dnterg dntete tinete a heat 8

roial eebsy PONTO] 08 CoH <4 dca lg BR Oe eR em el ORR et RR RO ARR OE AO RR eR Saye RO |

Si Basicdioimatay WHE? a ecieg 4 Slacks 4h soba Fk oct-g 4 testes Fd ston deg aen deg. bk og bbs ba ds 10

736 ... Krishnapriya & Kumar

oolivphacsmilated™ vex 2.5082. h65-7 Noe? Se. Sha, BERG SEN See oe SETS SES 2 Se. él

9. Hyphae not inflated;

basidiospores 4-4.5 x 3-5 um, subovoid, slightly fuscous........... C. gollanii

10. Basidiomata 6.5-8 cm high; basidiospores 3.3-3.7 x 5-7.4 um,

ellipsoid, convex on one side; hyphae not inflated, 3-7 um diam..... C. cretacea

10. Basidiomata 3-10 cm high; basidiospores 4.8-6.2 x 3.1-3.9 um,

not ellipsoid, pyriform; hyphae 2-3 um diam ....................0. C. fragilis

11. Basidiomata 2-3 cm high; basidiospores 4-6 x 2.5-3.5 um,

pyritorm; hyphae 15-18 um diame whe whee whee dy wabeleny abele wat C. crosslandii

11. Basidiomata 12 cm high; basidiospores 7-8 x 2.8-6.3 um,

subeylindric:hypliae Lui diay, 2g 0 ght Fue s8 2 ate ek mee ern he C. fumosa

Acknowledgements

The authors thank the Editor-in-Chief Dr. Lorelei Norvell and Nomenclature

Editor Dr. Shaun Pennycook for editorial corrections, and Dr. Ivona Kautmanova

(Slovak National Museum, Slovakia) and Dr. Anton G. Shiryaev (Ural Division of

the Russian Academy of Science, Russia) for providing critical reviews and valuable

suggestions for improvement of the manuscript. The authors also thank the Chief

Conservator of Forests & Chief Wildlife Warden, Kerala, for permission for fieldwork

in the forest areas of Kerala.

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https://doi.org/10.1016/B978-0-12-372180-8.50042-1

MY COTAXON

October-December 2021—Volume 136, pp. 739-748

https://doi.org/10.5248/136.739

Aspicilia lixianensis, A. nivalis, and

A. pycnocarpa spp. nov. from China

LIN Liu’ & QIANG REN**

"College of Life Sciences, Shandong Normal University, Jinan 250014, China

? State Key Laboratory of Mycology, Institute of Microbiology,

Chinese Academy of Sciences, Beijing 100101, China

* CORRESPONDENCE TO: rendagiang@hotmail.com

ABSTRACT — As a result of our study of Aspicilia from China, three new lichen species—

A. lixianensis, A. nivalis, and A. pycnocarpa—are proposed and typified. Detailed descriptions,

comparisons with related species, morphological photographs, and phylogenetic analysis of

these taxa are provided.

Key worps — ITS, Megasporaceae, Pertusariales, secondary metabolites, taxonomy

Introduction

Aspicilia A. Massal. is a large genus containing ca. 200 species (Licking

& al. 2016). Since being established by Massalongo in 1852, its taxonomic

status has been very vague. It was long treated as a section of Lecanora

Ach. (Lecanoraceae) for a long time (Magnusson 1940, 1952; Zahlbruckner

1930), until Hafellner (1984) restored Aspicilia to genus rank. Based on their

three-gene phylogenetic analysis, Lumbsch & al. (2007) placed Aspicilia in

Megasporaceae.

The main characteristics of Aspicilia are thallus crustose, rimose to

areolate, prothallus present or not (if present, sometimes peripherally

radiate or zonate); apothecia lecanorine, usually immersed, disk usually

concave; epihymenium olive-green or dark brown and turning green with

HNO,; paraphyses usually moniliform or submoniliform; 8-spored asci;

740 ... Liu & Ren

TABLE 1. ITS sequences used in the phylogenetic analysis.

Newly generated sequences are in bold.

SPECIES GENBANK SPECIES GENBANK

Aspicilia blastidiata KX129963 Aspicilia rivulicola EU057922

Aspicilia cinerea MN906262 Aspicilia subepiglypta KY249608

Aspicilia dendroplaca HQ259260 Aspicilia subgeographica KY249612

Aspicilia dudinensis EU057906 Aspicilia subgoettweigensis KY249614

Aspicilia goettweigensis KX159293 Aspicilia supertegens EU057938

Aspicilia lixianensis MW174005 Aspicilia virginea HQ259271

Aspicilia mashiginensis EU057912 Aspicilia volcanica NRI158307

Aspicilia nivalis MW174002

Aspicilia permutata EU057920 Lobothallia crassimarginata JX476026

Aspicilia pycnocarpa MW174004 Lobothallia pruinosa JX476028

simple, colorless, and often ellipsoid ascospores (Owe-Larsson & al. 2007).

Aspicilia species, which grow primarily on siliceous or calcareous rocks, are

widely distributed throughout the world.

Forty species of Aspicilia have been reported in China (Hou & al. 2014,

Ismayil & al. 2015, Li & al. 2013, Magnusson 1940, Shavkat & al. 2019,

Zahlbruckner 1930), but many specimens in Chinese herbaria remain

unidentified. After examining several specimen loans, we identified three

Aspicilia species new to science, described in this paper.

Materials & methods

The specimens examined were deposited in Herbarium Mycologicum Academiae

Sciences-Lichenes, Chinese Academy of Sciences, Beijing, China (HMAS-L) and

the Plant Herbarium of Shandong Normal University, Jinan, China (SDNU).

DNA extraction, PCR amplification, sequencing

DNA was extracted from the specimens using CTAB method. The isolated DNA

was amplified using primers ITSIF (Gardes & Bruns 1993) and ITS4 (White & al.

1990). Each 25 pL sample contained 5.5 uL distilled H,O, 5 uL DNA dilution, 1 uL of

each primer, and 12.5 uL CWBio 0690 2xEs Taq MasterMix (Dye). The PCR mixture

underwent a 5-minute initial denaturation at 95 °C, five cycles comprising 30 seconds

at 95 °C + 30 seconds at 58 °C + 1 minute at 72 °C followed by 30-35 cycles with the

Aspicilia lixianensis, A. nivalis, A. pycnocarpa spp. nov. (China) ... 741

annealing temperature decreased from 58 to 56 °C and concluded with final 7-minute

extension at 72 °C (Sohrabi & al. 2013). PCR products were sequenced by BioSune

Biotech Co. Ltd. in Beijing.

Phylogenetic analysis

We aligned three newly obtained ITS sequences with 14 ITS sequences belonging to

Aspicilia and two Lobothallia sequences as the outgroup downloaded from GenBank

(TABLE 1) using ClustalW in MEGA7 and optimized manually. A phylogeny was

inferred by the Maximum Likelihood (ML) method based on the General Time

Reversible model and Gamma Distributed in MEGA7 (Kumar & al. 2016).

Morphology and chemistry

We examined the morphology using a stereo microscope and a light microscope.

The lichen products were detected by color reaction tests (CT) and thin-layer

chromatography (TLC) using solvent systems A, B, and C (Elix 2014). The CT

97 KY249612 Aspicilia subgeographica

50 NR158307 Aspicilia volcanica

8 KX159293 Aspicilia goettweigensis

eal KY249614 Aspicilia subgoettweigensis

KX129963 Aspicilia blastidiata

97 58 KY249608 Aspicilia subepiglypta

78 MN906262 Aspicilia cinerea

MW174004 Aspicilia pycnocarpa

92 EU057906 Aspicilia dudinensis

85 EU057912 Aspicilia mashiginensis

= HQ259260 Aspicilia dendroplaca

EU057922 Aspicilia rivulicola

93 MW174005 Aspicilia lixianensis

EU057938 Aspicilia supertegens

MW174002 Aspicilia nivalis

EU057920 Aspicilia permutata

97 HQ259271 Aspicilia virginea

JX476026 Lobothallia crassimarginata

100 JX476028 Lobothallia pruinosa

0.020

Fic. 1. The most parsimonious ML tree generated from ITS sequences showing the phylogenetic

placement of Aspicilia pycnocarpa, A. lixianensis, and A. nivalis (in bold) among 17 Aspicilia spp.

and two Lobothallia spp. (outgroup). Bootstrap probabilities (50-100%) are indicated at internal

branches.

742 ... Liu & Ren

reagents used were K (10% KOH in water), N (50% nitric acid), and IKI (Lugol’s

iodine solution).

ITS sequence analysis

Phylogenetic analysis produced a parsimonious tree with 50-100%

bootstrap support values (Fic. 1). All Aspicilia species clustered within

one monophyletic clade with the ITS sequence analysis supporting

Aspicilia pycnocarpa, A. lixianensis, and A. nivalis as separate taxa. Aspicilia

pycnocarpa appears to be closely related to Aspicilia dudinensis. Aspicilia

lixianensis and A. nivalis are on separate branches, and no close relationships

are indicated by the analysis.

Taxonomy

Aspicilia pycnocarpa Q. Ren & Lin Liu, sp. nov. FIG. 2

EN 570766

Differs from Aspicilia cinerea by its longer conidia and possession of stictic acid as its

major lichen product.

Type: China, Jilin Province, route from Changbai County to Tianchi south slope scenic

area, alt. 1300 m, on siliceous rock, 25 Jul. 2014, L. Hu 20141380 (Holotype, SDNU;

GenBank MW174004).

ErymMo.oey: from the Greek elements pycno- (crowded) + -carpa (fruit), referring to

its abundant apothecia.

THALLUS crustose, areolate to rimose, 2.5-4.5 cm diam., 0.1-0.3 mm thick,

pale brown to brown; areoles irregular, 0.2-1 mm diam., flat, continuous,

fissured, cracks deep; prothallus inconspicuous. Upper cortex 25-30 um

thick, epruinose. Algal layer 20-30(-37.5) um thick.

APOTHECIA prominent, abundant, aspicilioid, 1(-4) per areole,

0.1-0.7 mm diam.; disc brown to black, flat to slightly concave, epruinose,

or sometimes slightly pruinose; thalline margin inconspicuous. True exciple

to 37.5-62.5 um wide in lateral portion. Epihymenium olive-green to olive-

brown, K+ brown, N+ green; Hymenium hyaline, IKI+ blue, 87.5-112.5 um

high; paraphyses not separating in KOH, distinctly moniliform with 2-6

globular cells; hypothecium colorless, IKI+ blue, 50-75 um thick, with algae

below the hypothecium. Asci clavate, Aspicilia-type, 8-spored; ascospores

hyaline, simple, ellipsoid, (12.5-)17.5-21.25 x (8.75-)10-12.5 um.

Pycnidia black, round-shaped; conidia filiform, straight or slightly curved,

(13.75-)17.5-25(-27.5) um long.

SPOT TESTS— Upper cortex K+ yellow, sometimes turning red or orange;

medulla K+ yellow.

Aspicilia lixianensis, A. nivalis, A. pycnocarpa spp. nov. (China) ... 743

wo,

7 P< NS

FiG. 2. Aspicilia pycnocarpa (holotype, SDNU 20141380). A. Thallus with apothecia; B. Apothecial

section; C. Ascus with 8 ascospores; D. Ascospores: colorless, simple, ellipsoid; E. Filiform conidia.

Scale bars: A = 1 mm; B = 100 um; C-E = 20 um.

SECONDARY METABOLITES—Stictic acid (major), norstictic acid (minor)

and substictic acid (minor) (TLC).

EcoLoGcy—On siliceous rock, interspersed with other lichens, such as

Lecidea sp. Known only from the type locality.

ADDITIONAL SPECIMEN EXAMINED—CHINA. JILIN PROVINCE: route from Changbai

County to Tianchi south slope scenic area, alt. 1300 m, on siliceous rock, 25 Jul. 2014,

L. Hu 20141379 (SDNU).

COMMENTS—Aspicilia pycnocarpa is characterized by its thin flat thallus, flat

to slightly elevated thalline margin, abundant apothecia, medium ascospores,

and long conidia. The specimens in SDNU were previously identified as

A. cinerea (L.) K6érb. based on the presence of norstictic acid, but A. cinerea

has shorter conidia (11-16 um) and contains mainly norstictic acid (Owe-

Larsson & al. 2007). Aspicilia pycnocarpa is similar to A. pacifica Owe-Larss.

& A. Nordin, which is distinguished by its higher hymenium (130-220

um), larger ascospores (14-33 x 10-20 um), and shorter conidia (8-17 um)

(Owe-Larsson & al. 2007).

744 ... Liu & Ren

Fic. 3. Aspicilia lixianensis (holotype, HMAS-L 0139004). A. Thallus in moist environment;

B. Thallus with apothecia; C. Apothecial section; D. Ascus with 8 ascospores; E. Ascospores:

colorless, simple, ellipsoid; F. Filiform conidia. Scale bars: A = 2 mm; B = 0.5 mm; C = 100 um;

D = 20 um; E, F = 10 um.

In the ML-tree, A. pycnocarpa appears closely related to A. dudinensis

(H. Magn.) Oxner, which differs by its usually disjointed pale or dark gray

areoles, smaller ascospores (15-18 x 8.5-10 um), and 18-22 tm long conidia

(Magnusson 1952, as Lecanora dudinensis).

Aspicilia lixianensis Q. Ren & Lin Liu, sp. nov. Fic. 3

EN 570763

Differs from Aspicilia asiatica by its higher hymenium, from Aspicilia subconfluens by its

lower hypothecium, and from both by its siliceous substrate.

Type: China, Sichuan Province, Lixian County, roadside of Jiabigou, 31.621 31.6214°N

1028455°E, alt. 2950 m, on siliceous rock, 10 Aug. 2016, W.C. Wang, WWC-51

(Holotype, HMAS-L 0139004; GenBank MW174005).

ETyMo_oey: referring to its type locality, Lixian County in Sichuan Province.

THALLUS crustose, areolate to rimose, 0.5-1.5 cm diam., +0.3 mm thick, gray to

dark gray; areoles irregular, 0.2-1 mm diam., flat or slightly convex, continuous,

fissured, cracks deep; prothallus inconspicuous. Upper cortex 12.5-25 um

thick, densely pruinose. Algal layer 37.5-50 um thick.

APOTHECIA prominent, aspicilioid, usually solitary in the areoles,

0.25-0.7 mm diam.; disc black, flat to slightly convex, with dense white pruina

Aspicilia lixianensis, A. nivalis, A. pycnocarpa spp. nov. (China) ... 745

in center; thalline margin usually inconspicuous. True exciple inconspicuous.

Epihymenium olive-brown, K+ brown, N+ green; hymenium hyaline,

IKI+ blue, 125-167.5 um high; paraphyses separating in KOH, distinctly

moniliform with 4-6 globular cells; hypothecium colorless, IKI+ blue,

37.5-87.5 um thick, with algae below the hypothecium. Asci clavate,

Aspicilia-type, 8-spored; ascospores hyaline, simple, ellipsoid, (10—)12.5-

17.5(-20) x (6.25—-)8.75-12.5 um. Pycnidia black, stellate-shaped; conidia

filiform, straight or curved, (10—)15-20 um long.

SPOT TESTS— Upper cortex K+ pale yellow; medulla K+ yellow.

SECONDARY METABOLITES—Stictic acid (major), substictic acid (minor)

(TLC).

EcoLtocy—On siliceous rock, growing with A. cinerea. Known only from

the type locality.

COMMENTS—Aspicilia lixianensis is characterized by its dark gray upper

surface with white pruina, black disks covered by densely white pruina,

medium ascospores and conidia, and the presence of stictic and substictic

acids. This specimen had few apothecia and abundant pycnidia and was

surrounded by A. cinerea.

Morphologically, A. lixianensis resembles A. asiatica (H. Magn.) Yoshim.

and A. subconfluens (H. Magn.) J.C. Wei, but those two species grow on

slightly calciferous rocks in arid and semi-arid areas. Additionally, A. asiatica

has lower hymenium (85-100 um) (Magnusson 1940, as Lecanora asiatica)

while the hypothecium of A. subconfluens is usually higher (150-200 um;

Magnusson 1940, as Lecanora subconfluens).

Aspicilia nivalis Q. Ren & Lin Liu, sp. nov. Fia. 4

EN 570764

Differs from Aspicilia decorticata by its white thallus, its higher hymenium, and its larger

ascospores.

Type: China, Xizang, Jiangda County, Xuejila Mountain, 31.445°N 98.0547E, alt. 4300

m, on sandstone, 10 Aug. 2016, W.C. Wang, WWC-148 (Holotype, HMAS-L 0139012;

GenBank MW174002).

ErymMo.oey: from nivalis, referring to the rough snow-white thallus and high elevation

snow-covered (in winter) type locality on Xuejila Mountain.

THALLUS crustose, areolate, 1-3.5 cm diam., +0.5 mm thick, white; areoles

irregular, 0.5-1.5 mm diam., flat, continuous, rough, slightly to deeply

fissured; prothallus inconspicuous. Upper cortex 25-50 um thick, powdery.

Algal layer 50-62.5 um thick.

746 ... Liu & Ren

APOTHECIA prominent, aspicilioid, 1(-3) per areole, 0.2-0.6 mm

diam.; disc black, slightly concave, with dense white pruina; thalline

margin conspicuous and flat, usually forming a black rim. True exciple

inconspicuous. Epihymenium dark brown, K+ brown, N+ green; hymenium

hyaline, IKI+ blue, 125-150 um high; paraphyses separating in KOH,

distinctly moniliform with 4-6 globular cells; hypothecium colorless, IKI+

blue, 25-45 um thick, without algae below the hypothecium. Asci clavate,

Aspicilia-type, 8-spored; ascospores hyaline, simple, ellipsoid, 18.75-30

x 10-15 um. Pycnidia black, round-shaped; conidia filiform, straight or

curved, 17.5-25 um long.

SPOT TESTS— Upper cortex K-; medulla K-.

SECONDARY METABOLITE—Substictic acid (TLC).

EcoLtoGy—On red-brown sandstone. Known only from the type locality.

CoMMENTS—Aspicilia nivalis is characterized by its continuous and

extremely rough thallus with powdery surface, large ascospores, and long

conidia. It is similar to A. decorticata (H. Magn.) J.C. Wei, which differs

by having a pale yellowish thallus, lower hymenium (85-100 um), smaller

ascospores (12-15 x 7.5-8.5 um), and calciferous rock substrate (Magnusson

1940, as Lecanora decorticata). The new species is also similar to A. persica

(Mull. Arg.) Sohrabi, which differs by growth on siliceous rock, densely

pruinose upper cortex, shorter ascospores (17.5-22.5 um), and different

secondary metabolites (stictic acid, +norstictic acid) (Li & al. 2013). Aspicilia

nivalis also resembles A. albocretacea (Zahlbr.) J.C. Wei, separated by its

continuous thallus with few cracks, epruinose discs, and shorter ascospores

(19-23 um; Zahlbruckner 1930, as Lecanora albocretacea).

Acknowledgements

Thanks to the curators of HMAS-L and SDNU for loan of specimens. We are

very grateful to Dr. Lulu Zhang (Institute of Environment and Ecology, Shandong

Normal University, China) and Dr. Xin Zhao (College of Life Sciences, Liaocheng

University, China) for reviewing the manuscript. This study was financially

supported by the National Natural Science Foundation of China (32070011).

Literature cited

Elix JA. 2014. A catalogue of standardized chromatographic data and biosynthetic relationships

for lichen substances. 3rd ed. Published by the Author, Canberra.

Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes -

application for the identification of mycorrhizae and rusts. Molecular Ecology 2: 113-118.

Hafellner J. 1984. Studien in Richtung einer naturlicheren Gliederung der Sammelfamilien

Lecanoraceae und Lecideaceae. Beihefte zur Nova Hedwigia 79: 241-371.

Aspicilia lixianensis, A. nivalis, A. pycnocarpa spp. nov. (China) ... 747

Fic. 4. Aspicilia nivalis (holotype, HMAS-L 0139012). A. Thallus on sandstone; B. Thallus with

apothecia; C. Apothecial section; D. Ascospores: colorless, simple, ellipsoid; E. Filiform conidia.

Scale bars: A = 1 mm; B = 0.5 mm; C = 100 um; D = 10 um; E = 20 um.

Hou Xj, Li SX, Ren Q. 2014. Three new records of aspicilioid lichens from China. Telopea 16:

159-163. https://doi.org/10.7751/telopea20147957

Ismayil G, Abbas A, Guo SY. 2015. Aspicilia volcanica, a new saxicolous lichen from Northeast

China. Mycotaxon 130: 543-548. https://doi.org/10.5248/130.543

Kumar S, Stecher G, Tamura K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis

version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870-1874.

https://doi.org/10.1093/molbev/msw054

Li SX, Kou XR, Ren Q. 2013. New records of Aspicilia species from China. Mycotaxon 126:

91-96. http://dx.doi.org/10.5248/126.91

Liicking R, Hodkinson BP, Leavitt SD. 2016. The 2016 classification of lichenized fungi in the

Ascomycota and Basidiomycota - approaching one thousand genera. Bryologist 119(4):

361-416. https://doi.org/10.1639/0007-2745-119.4.361

Lumbsch HT, Schmitt I, Licking R, Wiklund E, Wedin M. 2007. The phylogenetic placement

of Ostropales within Lecanoromycetes (Ascomycota) revisited. Mycological Research 111:

257-267. https://doi.org/10.1016/j.mycres.2007.01.006

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168 p.

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Flora of the Greater Sonoran Desert Region, vol. 3. Arizona State University, Tempe.

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Shavkat S, Ismayil G, Abbas A. 2019. Three new records of Aspicilia species from south slope

of Altai Mountains in Xinjiang. Acta Botanica Boreali-Occidentalia Sinica 39(1): 185-190.

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MYCOTAXON

October-December 2021— Volume 136, pp. 749-753

https://doi.org/10.5248/136.749

Architrypethelium barrerae sp. nov.

from a cloud forest in Veracruz, Mexico

JORGE GUZMAN-GUILLERMO* & REGULO CARLOS LLARENA- HERNANDEZ

Facultad de Ciencias Bioldgicas y Agropecuarias, Universidad Veracruzana,

Periuela, Amatlan de los Reyes, Veracruz, 94945 México

*CORRESPONDENCE TO: qwerty.guzmi@gmail.com

ABSTRACT—A new species of corticolous microlichen, Architrypethelium barrerae, is

described from the cloud forest in the ecological park ‘El Haya’ in Xalapa, Veracruz, Mexico.

It is characterized by two-spored asci, cortex UV+ yellow, and an internal K+ pigment in its

pseudostromata. Photographs of macro- and microscopic structures are presented.

Key worps—tropical lichens, Trypetheliales, Trypetheliaceae

Introduction

Cloud forests contain the highest vascular plant diversity in Mexico

(Rzedowski 1996). Despite the rich documentation of the vascular flora

there, lichen diversity from these, as with other tropical regions in Mexico, is

poorly known. Although some monographs and synopses have improved our

understanding of tropical lichens for some genera (e.g., Ahti 2000, Kurokawa

1962, Liicking 2008), overall little is known about the entire tropical lichen

flora. New species have been described from tropical regions (e.g., Cordova-

Chavez & al. 2014, Guzman-Guillermo & al. 2019, Hodkinson & al. 2014) ata

rather fast rate considering the paucity of specimens from such regions. This

highlights the potential for a great number of overlooked species yet to be

described.

Tropical Mexican lichens such as Trypetheliaceae are very poorly known, and

few reports have been published. This family comprises 16 mostly corticolous

genera with perithecioid ascomata, with or without stromatic tissues, and

750 ... Guzman-Guillermo & Llarena Hernandez

sometimes with xanthones and anthraquinones (Aptroot & Liicking 2016).

Within this family, Architrypethelium Aptroot contains perithecioid crustose

lichens similar to Astrothelium Eschw., but with large (>100 um), brown,

often 3-septate ascospores (Aptroot 1991). To date, this genus includes eight

accepted species, of which six are known to occur in the neotropics (Aptroot

& al. 2008, Aptroot & Licking 2016, Flakus & al. 2016, Licking & al. 2016),

one is paleotropical (Luangsuphabool & al. 2018), and one has a pantropical

distribution (Aptroot & Liicking 2016).

As part of a larger project, a survey was carried out to collect microlichens in

the megadiverse region in the center of Veracruz, a state of Mexico. We collected

corticolous microlichens in tropical vegetation, such as the cloud forest in

the ecological park ‘El Haya’ in Xalapa city. One result of our efforts was the

discovery of a previously unknown Architrypethelium, which we propose as the

new species A. barrerae.

Materials & methods

The material studied is preserved in the mycology collections in the herbarium of

Universidad Veracruzana, Xalapa, Veracruz, Mexico (XALU) and the herbarium of

Escuela Nacional de Ciencias Biolédgicas, Mexico City, Mexico (ENCB). Specimens

were collected in El Haya ecological park in Xalapa, Veracruz, México, located at

19.5181°N 96.9436°W and 1300 m a.s.l. Terminology is based on Aptroot & al.

(2008) and Aptroot & Licking (2016), and morphological analysis was conducted

by conventional lichenological techniques following Brodo & al. (2001). Sections for

anatomical examination were made by hand under a Zeiss Stemi Dv4 stereoscope

and microscopical measurements were taken in Lugol's solution using a Zeiss Primo

Star microscope. Thin layer chromatography (TLC) was performed using solvent C

following conventional chromatography techniques described in Elix (2014). Spot

tests used were C (NaClO) and K (KOH 5%). To test hymenium amyloidity, sections

of ascomata were mounted with I (Lugol's solution) and resultant coloration was

observed.

Taxonomy

Architrypethelium barrerae Guzm.-Guill. & Llar.-Hern., sp. nov. Figs 1, 2

MB 837958

Differs from Architrypethelium hyalinum by its larger ascospores, its two-spored asci,

and K+ internal pigment in its pseudostromata.

Type: México, Veracruz. Municipio de Xalapa, Parque Ecoldgico ‘El Haya, 19.5181°N

96.9436°W. 3 October 2020, Guzman-Guillermo 1968 (Holotype, XALU 25326).

Erymo.tocy: dedicated to Dr. Clementina Barrera-Bernal who supported the

lichenological studies by the first author.

Architrypethelium barrerae sp. nov. (Mexico) ... 751

Fic. 1. Architrypethelium barrerae (holotype, XALU 25326): A, B. Immature ascospores;

C. Mature ascospore; D. Ascus with two spores. Scale bar: A-C = 20 um; D = 40 um.

THALLUS corticate, green to yellowish, smooth to uneven. ASCOMATA

trypethelioid, with apical ostioles, 0-7-1-2 mm diam., erumpent to prominent,

covered by thallus except for the ostiolar area. Ascomata internally with a yellow

pigment. Wall fully carbonized and K+ olive green, apical and lateral portions

thick, base thin and touching the substrate. Hamathecium not inspersed. Asc1

cylindrical, longer than 300 um. Ascosporgs two per ascus, 160-200 x 50-75

um, oblong-ellipsoid, triple-septate, outer lumina much smaller than inner

lumina, lumina rounded in the corners, brown when free from the ascomata.

CHEMISTRY—Cortex UV+ yellow, thallus K-, yellow pigment K+ yellow to

dark red. TLC: lichexanthone and an unidentified anthraquinone.

ECOLOGY & DISTRIBUTION—Corticolous in open areas, in cloud forest at

1300 m a.s.l. Sometimes covered with moss. Known only from the type locality

in the El Haya ecological park in Xalapa city, Veracruz, Mexico.

752 ...Guzman-Guillermo & Llarena Hernandez

ComMMENtTS—Architrypethelium barrerae is morphologically similar to the

other Architrypethelium species but contains lichexanthone in its cortex.

Within the genus, only A. hyalinum Aptroot is known to have this chemistry,

but its spores are shorter (160-200 x 30-50 um) and its asci are 4-8-spored

(Aptroot & Licking 2016). Another characteristic only found in A. barrerae

is the pseudostromata with a conspicuous internal yellow pigment which is

K+ yellow to dark red.

ADDITIONAL MATERIAL STUDIED: MEXICO, VERACRUZ. Municipio de Xalapa, Parque

Ecoldégico ‘El Haya, 19.5181°N 96.9436°W, 1300 m a.s.l., 3 October 2020, Guzman-

Guillermo 1969, 1970, 1971, 1972, 1973, 1974, 1975, 1976, 1977 (XALU).

Acknowledgments.

We thank Dr. André Aptroot (Universidad Federal de Mato Grosso do Sul, Brazil)

for his review of this work and for the help provided in the study of microlichens in

Veracruz and Dr. Curtis Bjork (University of British Columbia, Vancouver, Canada)

for his presubmission manuscript review. We also thank the Environmental Quality

Laboratory and the herbarium XALU in the Facultad de Biologia of Universidad

Veracruzana for providing the resources needed to analyze the lichen samples studied

here and Subdireccién de Recursos Naturales y Cambio Climatico of Municipio de

Xalapa for providing permission to collect lichen samples from El Haya Ecological

Park.

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MY COTAXON

October-December 2021— Volume 136, pp. 755-767

https://doi.org/10.5248/136.755

Flavodontia rosea gen. & sp. nov.

from southwestern China

Hu1 WANG?” & CHANG-LIN ZHAO’

"Key Laboratory for Forest Resources Conservation and Utilization

in the Southwest Mountains of China, Ministry of Education

? College of Biodiversity Conservation,

'2 Southwest Forestry University, Kunming 650224, PR. China

“ CORRESPONDENCE TO: fungichanglinz@163.com

ABSTRACT—A new white-rot corticioid wood-inhabiting fungal genus and _ species,

Flavodontia rosea, collected from subtropical Yunnan, China, is proposed based on

morphological and molecular evidence. Flavodontia is characterized by annual, resupinate

basidiomes with a pink hymenial surface, a monomitic hyphal system with generative hyphae

bearing simple septa, and ellipsoid basidiospores with thin hyaline smooth walls. Phylogenetic

analyses of ITS and ITS + LSU nuclear RNA gene regions showed that Flavodontia formed a

distinct, monophyletic lineage within a subclade that includes Flavodon and Irpex.

Key worps—Irpicaceae, taxonomy, wood-rotting fungi

Introduction

Polyporales, a now well accepted and strongly supported order of

Agaricomycetes, is one of the most intensively studied groups of fungi of special

interest to fungal ecologists and applied scientists (Hibbett & al. 2014; Justo &

al. 2017). The most recent DICTIONARY OF THE FuNGI (Kirk & al. 2008) notes

1589 genera and more than 30,000 species of basidiomycetes (nearly 32% of all

described fungal taxa; Dai & al. 2015), of which roughly 1800 described species

are included in Polyporales, accounting for only about 1.5% of all known fungal

species (Kirk & al. 2008). These taxa play a key role as wood-rotting fungi

in addition to their pathogenicity and potential applications in biomedical

756 ... Wang & Zhao

engineering and biodegradation (Dai & al. 2009, Levin & al. 2016, Bankole &

al. 2020).

Polyporales has been sampled extensively in phylogenetic studies using

ribosomal RNA (rRNA) genes (Hibbett & Vilgalys 1993, Hibbett & Donoghue

1995, Boidin & al. 1998, Larsson & al. 2004, Binder & al. 2005, Justo & al. 2017,

Huang & al. 2020). Justo & al. (2017) included 18 families in their analyses of

a 3-gene dataset (292 taxa), providing a phylogenetic overview of Polyporales

as well as a framework for further taxonomic study. Among these, Irpicaceae

Spirin & Zmitr. encompassed a great variation of basidiome and hymenophore

types, with typical polypore and corticioid morphologies intermixing with

each other. A survey of morphological, anatomical, physiological, and genetic

traits supported 12 genera in this family.

During investigations of wood-inhabiting fungi in southern China, an

unknown taxon was found that could not be assigned to any described genus.

Here we expand samplings from other studies to examine the taxonomy and

phylogeny of a new genus within the Irpicaceae based on the internal transcribed

spacer (ITS) regions and the large subunit nrRNA gene (nLSU) sequences.

Materials & methods

The specimens are deposited at the herbarium of Southwest Forestry University,

Kunming, P.R. China (SWFC). Macroscopical descriptions are based on field notes

of fresh material. Colour terms follow Petersen (1996). Dried specimens were

examined microscopically under a light microscope following Dai (2012). The

following abbreviations are used: KOH = 5% potassium hydroxide, CB = Cotton

Blue, CB- = acyanophilous, IKI = Melzer’s reagent, IKI- = both inamyloid and

nondextrinoid, L = mean spore length (arithmetic average for all spores), W =

mean spore width (arithmetic average for all spores), Q = variation in the L/W

ratios between the specimens studied, n (a/b) = number of spores (a) measured

from given number of specimens (b).

A Magen Biotech HiPure Fungal DNA Mini Kit IT (Co., Ltd, Guangzhou,

Guangdong, P.R. China) was used to extract genomic DNA from dried specimens

according to the manufacturer's instructions with some modifications. A small

piece (ca. 30 mg) of dried fungal material was ground to powder with liquid

nitrogen, transferred to a 1.5 mL centrifuge tube, suspended in 0.4 mL of lysis

buffer, and incubated at 65°C in a water bath for 60 min. After 0.4 mL phenol-

chloroform (24:1) was added to each tube, the suspension was shaken vigorously.

After centrifugation at 13,000 rpm for 30 s, 0.3 mL of supernatant was transferred

to a new tube and mixed with 0.45 mL of binding buffer. The mixture was then

transferred to an adsorbing column (AC) for centrifugation again for 30 s. Then,

Flavodontia rosea gen. & sp. nov. (China) ...

TABLE 1. Sequences selected for ITS and nLSU phylogenetic analyses

SPECIES NAME

Byssomerulius corium

Ceriporia reticulata

C. viridans

Emmia lacerata

E. latemarginata

Flavodon flavus

Flavodontia rosea

Gloeoporus dichrous

G. pannocinctus

G. thelephoroides

Hydnopolyporus

fimbriatus

Irpex lacteus

Leptoporus mollis

Meruliopsis

leptocystidiata

M. taxicola

Phanerochaete rhodella

P. sordida

Trametopsis aborigena

T. brasiliensis

T. cervina

SPECIMEN NO.

FP-102382

Wu 1708-327

CBS 462.50

RLG-11354-Sp

Cui 8012

TNM: GC 1708-211

FP-55521T

Dai 7165

CBS 436.48

TNM: WHC 1381

LE295997

CLZhao 18491 [T]

CLZhao 18489

CBS 446.50

Dai 16370A

FCUG 2019

CBS 291.71

BZ-2896

CBS 384.51

FD-9

CBS 431.48

RLG7163

TJV-93-174T

TNM: Wu 1708-15

TNM: Wu 1708-43

CBS 455.48

BU061013-38

FD-18

FD-241

Robledo 1236

Robledo 1238

Meijer 3637

PRM900574

GENBANK ACCESSION NO.

ITS

KP135007

LC427007

MH856710

KP135041

KC182774

LC427027

KP135024

KY131834

MH856427

LC427029

KF856505

MW377575

MW377574

MH856705

KU360399

AF141612

MH860130

MG572757

MH856910

KP135026

MH856423

KY948794

KY948795

LC427012

LC427013

MH856432

MG572756

KP135187

KP135136

KY655336

KY655337

JN710510

AY684175

nLSU

KP135230

LC427031

MH868228

KP135204

LC427049

KP135202

KY131893

MH867973

LC427052

KF856510

MW377578

MW377577

MH868222

KU360406

AF141612

MH871903

MG572741

MH868432

KP135224

MH867969

EU402510

LC427032

LC427033

MH867978

MG572740

KP135258

KP135252

KY655338

KY655339

JN710510

AY855907

REFERENCES

Justo & al. 2017

Chen & al. 2020a

Vu & al. 2019

Justo & al. 2017

Jia & al. 2013

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Floudas & Hibbett

2015

Wu & al. 2017

Vu & al. 2019

Chen & al. 2020a

Zmitrovich &

Malysheva 2014

This study

Vu & al. 2019

Yuan & al. 2016

Vu & al. 2019

Jung & al. 2018

Vu & al. 2019

Justo & al. 2017

Vu & al. 2019

Justo & al. 2017

Chen & al. 2020a

Vu & al. 2019

Jung & al. 2018

Justo & al. 2017

Gomez-Montoya &

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Gomez-Montoya &

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Gomez-Montoya &

al. 2017

Tomsovsky & al.

2006

POT

758 ... Wang & Zhao

0.5 mL of inhibitor removal fluid was added in AC for a centrifugation at 12,000

rpm for 30 s. After washing twice with 0.5 mL of washing buffer, the AC was

transferred to a clean centrifuge tube, and 100 mL elution buffer was added to the

middle of adsorbed film to elute the genomic DNA. The ITS region was amplified

with primers ITS5 and ITS4 (White & al. 1990), and the nLSU region was amplified

with primers LROR and LR7 (http://www.biology.duke.edu/fungi/mycolab/

primers.htm). The ITS PCR protocols involved initial denaturation at 95°C for 3

min, 35 cycles at 94°C for 40 s + 58°C for 45 s + 72°C for 1 min, and a final 10-min

extension at 72°C. The nLSU PCR protocols involved initial denaturation at 94°C

for 1 min, followed by 35 cycles at 94°C for 30 s + 48°C for 1 min + 72°C for 1.5

min, and a final 10-min extension at 72°C. The PCR products were purified and

directly sequenced at Kunming Tsingke Biological Technology Limited Company.

All newly generated sequences were deposited in GenBank (TABLE 1).

Sequences were aligned in MAFFT 7 (http://mafft.cbrc.jp/alignment/server/)

using the “G-INS-I” strategy for nLSU and “E-INS-I” strategy for ITS + nLSU and

manually adjusted in BioEdit (Hall 1999). Alignment datasets were deposited in

TreeBase (submission ID 27155). Phanerochaete sordida and P. rhodella sequences

obtained from GenBank were used as outgroup to root trees following Justo & al.

(2017) in the ITS (Fic. 1) and ITS + nLSU (Fie. 2) phylogenetic trees.

Maximum parsimony analyses were applied to the ITS and ITS + nLSU dataset

sequences following Zhao & Wu (2017), and the tree was generated in PAUP* v.

4.0b10 (Swofford 2002). All characters were equally weighted with gaps treated

as missing data. Trees were inferred using the heuristic search option with TBR

branch swapping and 1000 random sequence additions. Max-trees were set to

5000, branches of zero length were collapsed and all parsimonious trees were saved.

Clade robustness was assessed using a bootstrap (BT) analysis with 1000 replicates

(Felsenstein 1985). Tree length (TL), consistency index (CI), retention index (RI),

rescaled consistency index (RC), and homoplasy index (HI) were calculated for

each Maximum Parsimonious Tree. The data matrix was also analyzed using

Maximum Likelihood (ML) approach with RAxML-HPC2 through the Cipres

Science Gateway (www.phylo.org; Miller & al. 2009). Branch support (BS) for ML

analysis was determined by 1000 bootstrap replicates.

MrModeltest 2.3 (Nylander 2004) was used to determine the best-fit evolution

model for each data set for Bayesian inference (BI). BI was calculated with MrBayes

3.1.2 with a general time reversible (GTR) model of DNA substitution and a

gamma distribution rate variation across sites (Ronquist & Huelsenbeck 2003).

Four Markov chains were run for 2 runs from random starting trees for 70,000

generations for ITS (Fic. 1), 200,000 generations for ITS + nLSU (Fic. 2) and

trees were sampled every 100 generations. The first one-fourth generations were

discarded as burn-in. A majority rule consensus tree of all remaining trees was

calculated. Branches were considered as significantly supported if they received

maximum likelihood bootstrap (BS) >70%, maximum parsimony bootstrap (BT)

>50%, or Bayesian posterior probability (BPP) >0.95.

Flavodontia rosea gen. & sp. nov. (China) ... 759

A tice 9 Allantoia . RS * « re oe A

A. Bydnoia 2 Broadly ovoid sto* “ee gr | yy »

A, Daedaloid to irpicoid 26 Cylindrical ES s >

Smooth OE Ellipsoid

- 9€. Narrowly elipsoia —Hoo1t00/.00 f: sa sane 431.48 A ¥ @ * *

iti Subcylindrical 78/53/0.98 | irpex lacteus FD-

M4 mane bata seangs| [-lootsn.0s ;-Flavodonflavus LE298997 A .") e * *

r Flavodonflavus TNM:WHC 1381

| |_jFlavodontiarosea CLZhao 18489 A ¥ es * *

@ Clamp connections 92/84/1.00 | 1°0/100.00 | FYavodontiarosea CLZhao 18491 T

@ Simple septate [ | ; Emmia latemarginata Dai 7165 A ) e@ * *

74994! emmia latemarginata CBS 436.48

Ye Narrowly clavate to cylindrical 190 100 100, L} — Ugmmiatacerata FP-5$52\T A yy @ * x

ke Fusiform — Hydnopolyporus fimbriatus CBS 384.51 A y es * *

fe Cylindrical —r.. Byssomeruliuscorium Wu 1708-327 A y e * x

afr Hans — 100/100/1.00 Byssomerulius corium FP-102382

1011001 00 | Meruliopsis taxicola CBS 455.48 A y e * x

Meruliopsis taxicola BU061013-38

“ee | Meruliopsis leptocystidiata TNM:Wu 1708-15 A y [ ) * *

1001100/1.00| \feruliopsis leptocystidiata TNM:Wu 1708-43

| e 100/100/1.00 [——Leptoporusmollis RLG7163 A y t } * x

LL Leptoporus mollis TIV-93-174T

100/100/1.00 (- Ceriporiareticulata CBS 462.50 A y @ * x

——#8-___ | Ceriporiareticulata RLG-11354

100/100/1.00 Ceriporiaviridans TNM:GC 1708-211 A y e * >" 4

ca Ceriporiaviridans Cui 8012

a | 1001880 99) Trametopsisaborigena Robledo 1238 A yp e k x

78/99/- || Trametopsisaborigena Robledo 1236

100/1001.00_| | Tyametopsiscervina PRM900S74 A .") e * x

— Trametopsis brasiliensis Meijer 3637 A w~ & * x

90/100/1,00 Gloeoporus dichrous CBS 446.50 A y e@ * x

iss. ~ Gloeoporus dichrous Dai 16370A

insti bs | | Gloeoporus pannocinctus CBS 291.71 A y t ) * *

ee eo) 100/100/1.00 | Gloeoporus pannocinctus FCUG 2019

Gloeoporus thelephoroides BZ-2896 A y e * x

Phanerochaeterhodella FD-18

Phanerochaete sordida FD-241

Fic. 1. Maximum Parsimony strict consensus tree illustrating the phylogeny of Flavodontia rosea

and related species in Irpicaceae based on ITS sequences. Branches are labelled with maximum

likelihood bootstrap >75%, parsimony bootstrap >50%, and Bayesian posterior probability >0.95.

Phylogenetic results

The ITS dataset included sequences from 33 fungal specimens

representing 20 taxa and with an aligned length of 666 characters, of which

358 characters were constant, 36 variable and parsimony-uninformative, and

272 parsimony-informative. Maximum parsimony analysis yielded 3 equally

parsimonious trees (TL = 927, CI = 0.537, HI = 0.463, RI = 0.731, RC =

0.393). Best model for ITS estimated and applied in the Bayesian analysis:

GTR+I4+G, Iset nst = 6, rates = invgamma; prset statefreqpr = dirichlet

(1,1,1,1). Bayesian analysis produced a similar topology with an average

standard deviation of split frequencies = 0.009412.

The phylogenetic tree (Fic. 1) inferred from ITS sequences obtained

from related genera in Irpicaceae demonstrated that the new genus formed

a monomitic lineage and then grouped with a clade including Flavodon

Ryvarden and Irpex Fr.

The ITS + nLSU dataset (Fic. 2) included sequences from 33 fungal

specimens, also representing 20 species. The dataset had an aligned length

of 1702 characters, of which 1120 characters were constant, 146 variable

and parsimony-uninformative, and 436 parsimony-informative. Maximum

760 ...

Wang & Zhao

i at

A Pore * She a2 ¥ ashe Rn oe & ot”

A. Hydnoid 2 Broadly ovoid oi wes a Oy Loe

A, Daedaloid to irpicoid 26 Cylindrical M vi j ig

A. Smooth 26 Ellipsoia (—ioorton.00 Irpex lacteus CBS 431.48 A oe e@ * b 4

9 Narrowly ellipsoid Irpexlacteus FD-9

@ Monomitic 2 Subcylindrical , l | 100/100/1.00 _[— Flavedontiarosea CLZhao 18491 T A y e@ * 4

| Bitohawise re Flavodonflavus LE295997 A gy e * x

Cline cities 100/10011.00| _ Flavodonflavus TNM:WHC 1381

@ Sine septs 104/100/1.00 1 Emmialatemarginata CBS 436.48 A ¥ e * *

-169!- |< Emmia latemarginata Dai 7165

100 96'L.09 ISL —— Emmia lacerata FP-55521T A y e@ * x

Ye Narrowly clavate to cylindrical a3 Hydnopolyporus fimbriatus CBS 384.51 A w e * x

We Fusiform 100/100/1.00; Byssomerulius corium Wu 1708-327 A y gS * b 4

ke Cylindrical SH821.00 Byssomerulius corium FP-102382

ye None 97/7210.99 | Trametopsis aborigena Robledo 1238 A wo e@ * > 4

93/98/0.99 |! Trametopsis aborigena Robledo 1236

00/1 00/1.90 Trametopsis cervina PRM900574 A y e * x

— Trametopsis brasiliensis Meijer 3637 A J e * x

BL i/8 100/100/1.00 [—— Ceriporia viridans Cui 8012 A y @ * D4

i CLL Ceriporia viridans TNM:GC 1708-211

[| 100/100/1.00 r— Ceriporia reticulata CBS 462.50 A y fon * b 4

r ae 09 Ceriporiareticulata RLG-11354

100/10011.00 (— Leptoporus mollis RLG7163 A ey e * *

L_ \— Leptoporus mollis TIV-93-174T

| 100/100/1.00 ¢ Meruliopsis leptocystidiata TNM:Wu 1708-15 A yy t ) * x

| ' Meruliopsis leptocystidiata TNM:Wu 1708-43

97/88/1.00 so0/oan,00 Meruliopsis taxicola CBS 455.48 A J e@ * x

Meruliopsis taxicola BU061013-38

100/100/1.00 i — Gloeoporusdichrous Dai 16370A A y e@ * *

Gloeoporus dichrous CBS 446.50

Soak 100/100/1.00_-— Gloeoporus pannocinctus CBS 291.71 A y e * > 4

_ Gloeoporus pannocinctus FCUG 2019

Gloeoporus thelephoroides BZ-2896 A y e@ * x

.-——— Phanerochaete rhodella FD-18

Phanerochaete sordida FD-241

Fic. 2. Maximum Parsimony strict consensus tree illustrating the phylogeny of Flavodontia

rosea and related species in Irpicaceae based on ITS + nLSU sequences. Branches are labeled

with maximum likelihood bootstrap >75%, parsimony bootstrap >50%, and Bayesian posterior

probability >0.95.

parsimony analysis yielded 2 equally parsimonious trees (TL = 1382, Cl

= 0.587, HI = 0.413, RI = 0.754, RC = 0.442). Best model for the nLSU

dataset estimated and applied in the Bayesian analysis: GIR+I+G, lset nst

= 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). Bayesian and

ML analyses produced a similar topology as MP analysis, with an average

standard deviation of split frequencies = 0.009828.

The additional phylogeny (Fic. 2) inferred from ITS + nLSU sequences

grouped the new species with the related species Flavodon flavus (Klotzsch)

Ryvarden and Irpex lacteus (Fr.) Fr.

Taxonomy

Flavodontia C.L. Zhao, gen. nov.

MB 838322

Differs from Flavodon by its resupinate basidiomata, smooth hymenial surface,

monomitic hyphal system, and ellipsoid basidiospores.

TYPE sPEcIEs: Flavodontia rosea C.L. Zhao

ErymMo oey: Flavodontia (Lat.): referring to the closely related genus, Flavodon.

Flavodontia rosea gen. & sp. nov. (China) ... 761

Fic. 3. Flavodontia rosea (holotype, SWFC 018491).

Basidiomata. Scale bars: A = 0.5 cm; B = 1 mm.

BASIDIOMATA annual, resupinate, smooth hymenial surface, fragile to rigid

upon drying. Hyphal system monomitic; generative hyphae bearing simple

septa, IKI-, CB-; tissues unchanged in KOH. Cystidia and cystidioles absent.

762 ... Wang & Zhao

Basidia clavate. Basidiospores ellipsoid, hyaline, thin-walled, smooth, IKI-,

CB-.

TYPE OF ROT: white.

Flavodontia rosea C.L. Zhao, sp. nov. Fics 3, 4

MB 838323

Differs from Flavodon flavus by its resupinate basidiomata with buff to slightly rose to

rose hymenial surface, its monomitic hyphal structure, and its smaller basidiospores.

Type: China. Yunnan Province: Honghe, Pingbian County, Daweishan National Nature

Reserve, 22.93°N 103.68°E, alt. 1714 m, on an angiosperm trunk, 3 August 2019,

CLZhao 18491 (Holotype, SWFC 018491; GenBank MW377575, MW377578).

EryMo_oey: From rosea (Lat.), referring to the rose hymenial surface of the basidiome.

BASIDIOMATA annual, resupinate, coriaceous, without odour or taste when

fresh, becoming fragile to rigid upon drying, <8 cm long, 3 cm wide, 500-800

um thick. Hymenial surface smooth, buff to slightly rose to rose when fresh,

becoming rose-coloured upon drying. Subiculum very thin, 0.2 mm thick, buff

to slightly rose. Margin sterile, cream to buff, 2 mm wide.

HyPHAL STRUCTURE monomitic; generative hyphae with simple septa,

colourless, thin to thick-walled, rarely branched, 3-4.5 um diam., IKI-,

CB-; tissues unchanged in KOH.

HyYMENIUM cystidia and cystidioles absent; basidia clavate, 19-26.5 x 3-4

um, with four sterigmata and a basal simple septum; basidioles dominant,

in shape similar to basidia, but slightly smaller.

Basip1osPorEs ellipsoid, colourless, thin-walled, smooth, IKI-, CB-,

(3.1-)3.3-4.6(-4.8) x (2.5-)2.7-3.5 um, L = 3.99 um, W = 3.06 um, Q =

1.29-1.30 (n = 60/2).

TYPE OF ROT: white rot.

ADDITIONAL SPECIMEN EXAMINED: CHINA. YUNNAN PROVINCE. Honghe: Pingbian

County, Daweishan National Nature Reserve, on the angiosperm trunk, 3 August 2019,

CLZhao 18489 (SWFC 018489; GenBank MW377574, MW377577).

Comments: Flavodon flavus differs from Flavodontia rosea by pileate

basidiomata with a poroid hymenophore surface that becomes hydnoid to

irpicoid with time, a dimitic hyphal structure with thick-walled skeletal hyphae,

and larger basidiospores (5.5-6.5 x 3-4 um; Ryvarden 1973).

Discussion

Phylogenetic and morphological analyses strongly support Flavodontia

as an independent genus that clusters with related taxa in Irpicaceae.

Phylogenetically, Flavodontia is closely related to Flavodon and Irpex based

Flavodontia rosea gen. & sp. nov. (China) ... 763

Fic. 4. Flavodontia rosea (drawn from the holotype, SWFC 018491).

A. Basidiospores; B. Basidia and basidioles; C. Section of hymenium.

Scale bars: A = 5 um; B, C = 10 um.

on ITS and ITS + LSU nRNA gene analyses (Fics 1 & 2) in agreement

with multigene analyses by Justo & al. (2017). Flavodon is morphologically

distinguished by pileate basidiomata with poroid to irpicoid hymenophores

and encrusted cystidia (Ryvarden 1973), and Irpex differs from Flavodontia

by its poroid to hydnoid hymenophore with irregular/irpicoid pores and

dimitic hyphal system (Fries 1825, Bernicchia & Gorjoén 2010).

The consensus phylogram of the comprehensive taxon dataset from 12

agaricomycete orders from ITS, LSU, mtSSU, atp6, tefl, and rpb2 grouped

Flavodon, Irpex, and Trametopsis 'TomSovsky together in a clade labelled

“Byssomerulius family” (Miettinen & al. 2012). A subsequent phylogenetic

tree based on a 3-gene dataset and including more sequences and taxa in

Irpicaceae supported Emmia Zmitr. & al. as sister to Irpex and clustered with

Trametopsis (Justo & al. 2017). The relationships among different taxa in

Irpicaceae from both studies (Miettinen & al. 2012, Justo & al. 2017) differed

due to 3-gene and 6-gene datasets.

In the present study, the ITS (Fic. 1) and ITS + nLSU (Fic. 2) phylograms

are very similar, clustering four genera together in agreement with previous

studies and supporting Flavodontia as an independent genus. The different

764 ... Wang & Zhao

symbols and colors in Figs 1 & 2 show that the macromorphological

hymenophore characters are not consistent and regular across Flavodontia

and related genera. However, the micromorphological data (hymenial

surface, hyphal system, generative hyphae, and cystidial and spore shapes)

show that four closely related genera Emmia, Flavodon, Flavodontia, and Irpex

share the same character of simple septate generative hyphae, suggesting that

micromorphology is less affected by the environment than macromorphology

and thus more indicative of phylogenetic relationships.

Irpicioid fungi represent an extensively studied group of Polyporales

(Nunez & Ryvarden 2001, Bernicchia & Gorjén 2010, Dai 2012, Ryvarden &

Melo 2014, Dai & al. 2015), but the Chinese irpicioid fungus diversity is still

not well known, especially in subtropics and tropics where many new taxa,

including Flavodontia, have recently been described (Zhao & al. 2017, Shen

& al. 2018, Cui & al. 2019, Ma & Zhao 2019, Chen & al. 2020b, Huang & al.

2020). We anticipate that more undescribed irpicioid taxa will be discovered

throughout China after extensive collection combined with morphological

and molecular analyses.

Acknowledgements

The research was supported by the National Natural Science Foundation of

China (Project No. 32170004), Yunnan Fundamental Research Project (Grant No.

202001AS070043) and High-level Talents Program of Yunnan Province (YNQR-

QNRC-2018-111). The authors thank Mei-Ling Han (Langfang Normal University,

China) and Sana Jabeen (University of Education, Lahore, Pakistan) for helpful

comments during presubmission review.

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MYCOTAXON

October-December 2021— Volume 136, pp. 769-778

https://doi.org/10.5248/136.769

Pseudodeightoniella indica gen. and sp. nov.,

a hyphomycete from India

SANJEET KUMAR VERMA’, SANJAY YADAV', RAGHVENDRA SINGH’ ,

BALMUKUND CHAURASIA’, SHAMBHU KUMAR?

"Centre of Advanced Study in Botany, Institute of Science, Banaras Hindu University,

Varanasi, U.P, India 221005

?754-D, Ramjanaki Nagar, Mirzapur Pachpedwa, West Basharatpur,

Gorakhpur, U.P, India 273004

° Forest Pathology Department, KSCSTE-Kerala Forest Research Institute,

Peechi, Thrissur, Kerala, India 680653

“ CORRESPONDENCE TO: drsinghtaxon@gmail.com, singhr.bot@bhu.ac.in

ABSTRACT—An interesting new mitosporic foliicolous genus and _ species,

Pseudodeightoniella indica, discovered on living leaves of Medicago polymorpha

(Fabaceae) from Uttarakhand, India, is described and illustrated. Morphologically,

Pseudodeightoniella differs from the closely similar Deightoniella and Neodeightoniella in

producing well-developed erumpent stromata, highly undulate to spiral or helicoid and

densely fasciculate conidiophores with enlarged basal cells, and conidia lacking mucoid

caps. Recently published illustrations of conidia and conidiophores characterizing the

Deightoniella and Neodeightoniella type species are also provided.

KEY worDs—asexual fungi, biodiversity, plant disease, taxonomy

Introduction

Deightoniella S. Hughes was introduced with type species D. africana

S. Hughes (Hughes 1952). The main diagnostic features of Deightoniella are

mononematous brown conidiophores that are torsive or flexuous and typically

nodulated or annellated by the outer wall after each enteroblastic precurrent

proliferation of the conidiogenous cells and obclavate to obpyriform,

subhyaline to olivaceous brown, minute verruculose, transversely 1-2-septate

770... Verma & al.

conidia with the septum above the median and the apical cell prominently

tapering towards subobtuse apex (Hughes 1952, Klaubauf & al. 2014, Videira

& al. 2017). Multigenic analysis by Klaubauf & al. (2014) placed Deightoniella

in a new family, Pyriculariaceae.

Similar to Deightoniella is Neodeightoniella Crous & W.J. Swart, described

in 2013 with the type species N. phragmiticola Crous & WJ. Swart (Crous & al.

2013). Neodeightoniella differs from Deightoniella in producing less fasciculate,

straight (to slightly flexuous) sub-cylindrical conidiophores produced from

a poorly developed stromata of a few brown cells; terminal and integrated

conidiogenous cells; dark thickened scars; and 1-septate fusoid-ellipsoid

conidia with apical cells that are globose and with prominent mucoid cap

and funnel-shaped basal cells that are widest two thirds from the basal hilum

before tapering prominently to a truncate, darker and thickened hilum, hilum

with a central pore (Crous & al. 2013, Videira & al. 2017). Phylogenetically,

Neodeightoniella clusters in the Mycosphaerellaceae (Videira & al. 2017).

Fifteen Deightoniella and one Neodeightoniella species are currently accepted

(www.indexfungorum.org; accessed 15 August 2020).

During the past decade, several asexual foliicolous fungi have been

described from north India (Awasthi & al. 2016; Kumar & Singh 2015a,b,

2016; Kumar & al. 2012a,b, 2018; Kushwaha & al. 2020; Singh & Kamal 2011;

Singh & al. 2011, 2012, 2013a,b, 2014a,b, 2019), suggesting that such fungi

are highly diverse and in need of additional exploration and characterization.

During a systematic survey of conidial fungi in the forests of Nainital,

Uttarakhand, an interesting species was collected on young living leaves of

Medicago polymorpha. Morphologically, this unknown fungus somewhat

resembled Deightoniella and Neodeightoniella but was sufficiently distinct to

warrant our proposal of it as a new genus and species.

Materials & methods

In October 2018, infected leaves of Medicago polymorpha were collected near

Bhimtal of Nainital, Uttarakhand, during field survey, and the collected samples

were kept in sterilized polythene bag for transport to the laboratory where they were

processed according to standard techniques (Hawksworth 1974, Savile 1962). Slides

for microscopic examination were prepared by scraping specimens from infected

areas, hand sectioning, and mounted in clear lacto-phenol cotton blue mixture and

clear glycerin. The microphotographs of fungal propagules were taken using Magnus

MIPS CMOS camera attached to an Olympus CH20i-TR compound microscope.

The detailed morphological observations and Lucida drawings were conducted at

different magnifications (100x, 450x, 1000x), and 20 measurements were made of

each morphological feature. The holotype specimen was deposited in the Ajrekar

Pseudodeightoniella indica gen. and sp. nov. (India) ... 771

Fic. 1. Pseudodeightoniella indica on Medicago polymorpha. a. Host plant in natural habitat;

b. Symptom on upper surface of leaflets; c, d. Initial stages on infection on lower surfaces of

leaflets; e. Late stage of infection on lower surface of leaflet; f-h. Colonies of fungal pathogen at

high magnifications. Scales bars: b-e = 5 mm; f, g = 200 um; h = 100 um.

Mycological Herbarium, Agharkar Research Institute, Pune, India (AMH), and

an isotype is retained in the Mycological Herbarium of the Department of Botany,

Banaras Hindi University, Varanasi, India (MH-BHU).

772... Verma & al.

Taxonomy

Pseudodeightoniella S.K. Verma, Sanj. Yadav & Raghv. Singh, gen. nov.

MB 836237

Differs from Deightoniella and Neodeightoniella by its very well-developed erumpent

stromata, densely fasciculate and undulate to spiral or helicoid conidiophores with

swollen basal cell, and conidia without mucoid cap.

TyPE SPECIES: Pseudodeightoniella indica S.K. Verma & al.

Erymo.oey: In reference to partial morphological similarity to the genus Deightoniella.

Cotonies hypophyllous, scattered, dark brown to blackish brown.

MyceLium internal. Srromata well-developed, prosenchymatous to

pseudoparenchymatous, erumpent, mid- to dark brown. CONIDIOPHORES

macronematous, densely fasciculate, arising from stromata, sub-cylindrical,

straight to slightly flexuous, undulate to spiral/helicoid, unbranched, smooth

to finely roughened, mid- to dark brown, septate, bearing swollen tubular or

oval shaped basal cells that aggregate at the top of stromata. CONIDIOGENOUS

CELLS integrated and terminal, sub-cylindrical, mid brown to dark brown,

slightly lighter towards apex, finely roughened, enteroblastic, mono to

polytretic; scars terminal and lateral on conidiogenous cells, thickened and

darkened, slightly protruding, circular with central pore. Conidial secession

schizolytic. CONIDIA solitary, mid- to dark brown, surface smooth to finely

roughened, mostly obpyriform to cuneiform, slightly curved, apical cell

globose, basal cell more or less funnel shaped, aseptate to septate with septum

medially placed, tapering towards truncate hilum, hilum slightly thickened

and dark.

Pseudodeightoniella indica S.K. Verma, Sanj. Yadav &

Raghv. Singh, sp. nov. FIGS 1-4

MB 836238

Differs from Deightoniella africana by its septate conidiophores with larger basal cell and

smaller conidia without any kind of conical shaped apical cell and from Neodeightoniella

phragmiticola by its shorter, thinner conidiophores and smaller conidia lacking any

mucoid cap on apical cells.

Type: India, Uttarakhand, Nainital, Bhimtal, 29.3919°N 79.4542°E, on living leaves

of Medicago polymorpha L. (Fabaceae), 20 October 2018, coll. Sanjeet Kumar Verma

(Holotype, AMH 10239; isotype, MH-BHU 19).

Erymo oey: Derived from the name of country where the taxon was discovered.

INFECTION spots hypogenous, brown to dark blackish brown, velvety,

scattered, initially circular to subcircular but later becoming irregular,

0.2-1.1 mm in diam. CoLontes hypophyllous, scattered, dark brown to

Pseudodeightoniella indica gen. and sp. nov. (India) ... 773

Fig. 2. Pseudodeightoniella indica (holotype, AMH 10239). a. Initial stage of development

of stromata lined by basal swollen cells of conidiophores; b. Initial stage of development of

conidiophores from swollen basal cells; c, d. Fully developed stage of undulating conidiophores

in fascicle; e. Undulating nature of conidiophores; f-j. Different forms of conidiophores.

Scale bars: 10 um.

blackish brown. MyceLrtum immersed, medium brown. STROMATA well-

developed, prosenchymatous to pseudoparenchymatous, erumpent, mid

brown to dark brown, 30-70 x 50-90 um. CONIDIOPHORES macronematous,

77A. ... Verma & al.

Fic. 3. Pseudodeightoniella indica (holotype, AMH 10239). a-e. Undulating conidiophores

bearing basal swollen cells (green arrow for initial stage of development of conidiophores from

basal cell); f. Conidiophores with developing conidia; g-i. Enteroblastic nature of conidial

ontogeny (blue arrow); j, k. Conidiogenous cells with circular loci having central pore (red

arrows); l-r. Conidia. Scale bars: 10 um.

Pseudodeightoniella indica gen. and sp. nov. (India) ... 775

Fic. 4. Pseudodeightoniella indica (holotype, AMH 10239). a. Initial stage of development

of stromata lined by basal swollen cells of conidiophores; b. Initial stage of development of

conidiophores from swollen basal cells; c. Developed stage of undulating conidiophores in

fascicle; d. Initial developmental stage of conidiophores with basal cells; e. Different forms of

developed conidiophores with basal cells; f, g. Conidiogenous cells with loci; h. Enteroblastic

development of conidium; i. Conidia. Scale bars: 10 um.

densely fasciculate, arising from stromata, sub-cylindrical, straight to

slightly flexuous, undulate to spiral/helicoid, unbranched, smooth to

finely roughened, medium to dark brown, 0-2-septate, 20-50 x 2-6.5 um

(excluding basal cells), bearing swollen tubular or oval basal cells (15-26

x 7-14 um) that form aggregations (70-85 x 25-40 um) at the tops of

stromata. CONIDIOGENOUS CELLS integrated and terminal, sub-cylindrical,

mid brown to dark brown, slightly lighter towards apex, finely roughened,

enteroblastic, mono to polytretic, 6-16 x 3-6.5 um; scars terminal and lateral

TO 33:

Verma & al.

Fic. 5. Deightoniella africana: a. Conidia and conidiophores (Videira & al. 2017).

Neodeightoniella phragmiticola: b. Conidia and conidiophores (Crous & al. 2013).

Scale bars: 10 um.

on conidiogenous cells, thickened and dark, slightly protruding, circular

with central pore, 2.2-4.3 um diam. Conidial secession schizolytic. CONIDIA

solitary, medium to dark brown, surface smooth to finely roughened, mostly

obpyriform to cuneiform, slightly curved, 13.5-18.5 x 8.5-11.5 um, apical

cell globose, basal cell more or less funnel-shaped, aseptate to 1-septate,

septum medially placed, tapering towards truncate hilum, hilum slightly

thickened and darkened, 2.6-4.1 um wide.

CoMMENTS—Pseudodeightoniella shares a_ superficial morphological

similarity with Deightoniella (Hughes 1952, Klaubauf & al. 2014, Videira

& al. 2017) and Neodeightoniella (Crous & al. 2013, Videira & al. 2017)

due to the undulate nature of the conidiophores. Deightoniella, however,

is distinguished by its lack of stromata; solitary, torsive or flexuous

conidiophores; monotretic conidiogenous cells, and conidia that are

transversely 1-septate above the median and with the apical cell prominently

tapering towards the sub-obtuse apex; the conidiophore of Deightoniella

shows characteristic swellings along the conidiophore length resulting from

percurrent rejuvenation and conidiophore elongation (e.g., D. africana,

Fic. 5A). Pseudodeightoniella is distinct from Neodeightoniella, which has

Pseudodeightoniella indica gen. and sp. nov. (India) ... 777

very poorly developed stromata (composed of only a few cells), conidia with

globose apical cells bearing a prominent mucoid cap, and an absence of

percurrent rejuvenation and any swollen tubular or oval shaped basal cells

in the conidiophores (e.g., N. phragmiticola, Fic. 5B).

Acknowledgments

The authors express their sincere gratitude to Dr Patricia Oliveira Fiuza

(Universidade Federal do Rio Grande do Norte, Brazil) and Dr. Alisson Cardoso

Rodrigues da Cruz (Universidade Federal da Bahia, Brazil) for their critical reviews

of the manuscript and useful suggestions. The authors thank the curator of AMH,

ARI, Pune, for accepting the holotype material and providing an accession number.

Authors are also thankful to the Head of the Department of Botany, Institute of

Science, Banaras Hindu University, Varanasi, for providing laboratory facilities.

Dr. Lorelei Norvell’s editorial review and Dr. Shaun Pennycook’s nomenclature reviews

are greatly appreciated.

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foliicolous hyphomycetous genus from Uttar Pradesh, India. Kavaka 50: 64-68.

https://doi.org/10.5248/131.25

Kushwaha P, Singh R, Chaurasia B. 2020. Ramularia titarpaniensis—a new species of

ramularioid complex from central India. Phytotaxa 429 (4): 274-280.

https://doi.org/10.11646/phytotaxa.429.4.3

778 ... Verma & al.

Savile DBO. 1962. Collection and care of botanical specimens. Research Branch, Canadian

Department of Agriculture, Publication 1113. 124 p. https://doi.org/10.5962/bh1.title.53755

Singh R, Kamal. 2011. Two new species of Corynespora from northeastern Uttar Pradesh, India.

Mycotaxon 118: 123-129. https://doi.org/10.5248/118.123

Singh R, Kumar S, Kamal. 2011. Two new species of Passalora and Pseudocercospora from

northeastern Uttar Pradesh, India. Mycotaxon 117: 137-143. https://doi.org/10.5248/117.137

Singh R, Chaurasia B, Shukla K, Upadhyaya PP. 2012. Passalora aseptata, a new

cercosporoid fungus from northeastern Uttar Pradesh, India. Mycotaxon 120: 461-463.

https://doi.org/10.5248/120.461

Singh R, Kumar S, Singh A, Shukla K. 2013a. A new species of Parapyricularia from India and a key

to all species. Sydowia 65(2): 337-342.

Singh R, Kumar S, Saini DC, Upadhyaya PP, Kamal, Braun U. 2013b. Diversity of Passalora on

Ficus. Mycological Progress 12: 637-643. https://doi.org/10.1007/s11557-012-0870-6

Singh R, Singh A, Kumar S, Upadhyaya PP, Castafeda-Ruiz RE 2014a. Two new species of

Zasmidium from northeastern Uttar Pradesh, India. Nova Hedwigia 98(1-2): 257-263.

https://doi.org/10.1127/0029-5035/2013/0137

Singh A, Kharwar RN, Singh R, Kumar S. 2014b. A new species of Zasmidium (Mycosphaerellaceae)

from India. Sydowia 66 (2): 309-312.

Singh A, Singh NK, Singh PN, Singh R, Dubey NK. 2019. Additions to Ochroconis from India.

Phytotaxa 427(3): 186-199. https://doi.org/10.11646/phytotaxa.427.3.2

Videira SIR, Groenewald JZ, Nakashima C, Braun U, Barreto RW, de Wit PJGM, Crous

PW. 2017. Mycosphaerellaceae - chaos or clarity? Studies in Mycology 87: 257-421.

https://doi.org/10.1016/j.simyco.2017.09.003

MY COTAXON

October-December 2021— Volume 136, pp. 779-784

https://doi.org/10.5248/136.779

Lepra yunlingensis and L. taiwanensis spp. nov.

from China

JIARONG ZHANG’, XIANDONG XUE’, LIN Liu*”, XUYUN QIU’’, QIANG REN?

' College of Life Sciences, Shandong Normal University,

No. 88 East Wenhua Road, Jinan 250014, China

State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences,

No. 1 West Beichen Road, Beijing 100101, China

“CORRESPONDENCE TO: rendaqiang@hotmail.com

ABSTRACT—Two new species of Lepra are described from China: L. yunlingensis is

characterized by its 2-spored asci and the presence of norstictic and cryptostictic acids;

L. taiwanensis is characterized by its 1-spored asci and the presence of barbatic acid.

Key worps—lichenized fungi, Pertusaria, Pertusariaceae, Pertusariales, taxonomy

Introduction

Based on a combination of morphological and chemical characters,

Pertusaria DC. was divided into three subgenera (Archer 1993). The earlier

name Lepra Scop., rejected against the conserved Pertusaria, which could

accommodate the species of P subgen. Monomurata A.W. Archer (Archer

1993) or the “Variolaria”’-group (Schmitt & Lumbsch 2004) of Pertusaria s.lat.,

was resurrected by Hafellner & Tiirk (2016). That circumscription of the genus

Lepra is now accepted by other lichenologists (Lendemer & Harris 2017, Archer

& Elix 2018, Ren 2019).

Lepra is characterized by its crustose thallus, disciform apothecia, 1-2

(-rarely 8)-spored asci, large hyaline ascospores, and the presence of a diversity

of lichen substances, such as depsone, depsidones, B-orcinol m-depsides,

6-orcinol p-depsides, fatty acids, and lichexanthone (Archer & Elix 2018).

780 ... Zhang & al.

In the course of reexamining Lepra specimens in KUN, two new species of

Lepra were encountered that are described here.

Materials & methods

The specimens examined in this study were deposited in the Herbarium

at Kunming Institute of Botany, Chinese Academy of Sciences, Yunnan, China

(KUN).

Macromorphological characters were observed using a stereo microscope

and micromorphological characters were examined by hand-cut sections under

a light microscope. The lichen products were detected primarily by spot tests K =

a aqueous solution of 10% potassium hydroxide (KOH), C = a saturated aqueous

solution of calcium hypochlorite (Ca(ClO),), KC = K solution followed by C] and

thin-layer chromatography (TLC) with solvent systems A, B, and C (Elix 2014).

Taxonomy

Lepra yunlingensis Q. Ren, sp. nov. FIG. 1

FN 570769

Differs from Lepra wulingensis by its 2-spored asci and the absence of psoromic acid.

Type: China. Yunnan, Nanjian County, Lingbaoshan National Park, 24.7700°N

100.4961°E, alt. 2355 m, on bark, 29 Jun. 2015, X. Ye 15-47837 & W.C. Wang

(Holotype, KUN).

Erymo.oecy: The epithet refers to the name of the type locality, the Yunling Mountains.

THALLUS corticolous, crustose, verrucose to verruculose, thin, grayish green;

margin indistinct; prothallus not visible; isidia absent; soredia present,

granular, restricted to the verrucae. VERRUCAE sorediate, numerous, crowded

but rarely fused, c. 1.0 mm in diam. APOTHECIA 1 (less often 2-3) per verruca,

with pinkish discs densely covered with white pruina. AscosPoRES 2 per ascus,

uniseriate longitudinally, hyaline, 57-97 x 33-48 um; ascospore wall smooth.

PyYcNIDIA not observed.

CHEMIsTRY: Medulla K+ yellow turning red, C-, KC-. Containing norstictic

acid and cryptostictic acid.

REMARKS: Diagnostic characters for L. yunlingensis are a thin corticolous

thallus, granular soredia restricted to verrucae, pinkish discs covered with

white pruina, 2-spored asci, and the presence of norstictic and cryptostictic

acids. Morphologically, the new species is very similar to L. wulingensis

(Z.T. Zhao & Z.S. Sun) Q. Ren, which differs in its 8-spored asci and

presence of psoromic acid (Ren & al. 2009, as Pertusaria wulingensis).

Lepra taiwanensis & L. yunlingensis spp. nov. (China) ... 781

Fic. 1. Lepra yunlingensis (KUN - Ye 15-47837). A. Thallus with numerous verrucae. B. Detail of

verrucae, showing disks covered with white pruina. Scale bars = 1 mm.

Lepra bambusetorum (Zahlbr.) Q. Ren, L. composita (Zahlbr.) Q. Ren, and

L. trachythallina (Erichsen) Lendemer & R.C. Harris are three species known

from China that produce disciform apothecia and 2-spored asci. However, they

are separated from L. yunlingensis by the presence of different lichen substances:

782 ... Zhang & al.

salazinic acid in L. bambusetorum, protocetraric acid in L. composita, and

thamnolic acid in L. trachythallina (Ren 2019).

Lepra taiwanensis Q. Ren, sp. nov. FiG. 2

FN 570770

Differs from Lepra multipunctoides by its pinkish discs and the absence of

fumarprotocetraric acid.

Type: China. Taiwan, Nantou County, Mt. Hehuan, 24.1358°N 121.2872°E, alt. 3303 m,

on bark of Abies sp., 23 Sep. 2015, L.S. Wang 15-49266 & X.Y. Wang (Holotype, KUN).

EtymMo oey: The epithet refers to the name of the type locality, Taiwan.

THALLUS corticolous, crustose, thin, whitish to grayish; prothallus distinct,

whitish; isidia absent; soredia fine and restricted to the verrucae, white.

VERRUCAE abundant, rounded in outline, 1-1.5 mm in diam., punctiform to

hemispherical when young, pseudocyphella-like. APOTHECIA 1-4 per verruca,

with pinkish discs densely covered with white pruina. AscosporeEs 1 per

ascus, usually immature, hyaline, ellipsoid, 70-205 x 20-81 um; ascospore wall

smooth, c. 13 um thick. Pycnip1a not observed.

CHEMIsTRY: Medulla K-, C-, KC+ pink. Containing barbatic acid.

ADDITIONAL SPECIMEN EXAMINED—CHINA. Taiwan, Nantou County, Mt. Hehuan,

24.1358°N 121.2872°E, alt. 3303 m, on Abies bark, 23 Sep. 2015, L.S. Wang 15-49219 &

X.Y. Wang (KUN).

REMARKS: Diagnostic characters for L. taiwanensis are a thin whitish to

grayish corticolous thallus, fine soredia restricted to verrucae, pinkish discs

densely covered with white pruina, 1-spored asci, and the presence of barbatic

acid. Morphologically, the new species resembles L. multipunctoides (Dibben)

Lendemer & R.C. Harris and the abundantly collected species identified by

Chinese lichenologists as L. amara (Ach.) Hafellner, but L. multipunctoides

contains fumarprotocetraric acid and L. amara contains picrolichenic acid

(Ren 2019).

Barbatic acid is also known from two other Lepra species: L. barbatica (A.W.

Archer & Elix) I. Schmitt & al. and L. wirthii (Elix & A.W. Archer) I. Schmitt

& al. Lepra barbatica is a sterile, isidiate species occurring in Eastern Australia

and New Zealand (Archer 1997, Galloway 2007; as Pertusaria barbatica);

L. wirthii has black discs and larger ascospores (200-240 x 56-70 um) and

occurs in New Zealand (Archer & Elix 2013, as Pertusaria wirthii).

Acknowledgements

Thanks to the curator of KUN for loan of specimens, with special thanks to Mr.

Lisong Wang (KUN) and Dr. Xinyu Wang (KUN). The authors are grateful to Dr.

Lepra taiwanensis & L. yunlingensis spp. nov. (China) ... 783

Fic. 1. Lepra taiwanensis (KUN - Wang 15-49266). A. Thallus with substratum, showing numerous

verrucae. B. Young thallus with pseudocyphella-like verrucae. Scale bars = 5 mm.

Huajie Liu (College of Life Sciences, Hebei University, China) and Dr. Zefeng Jia

(College of Life Sciences, Liaocheng University, China) for reading and improving

the manuscript. This study was financially supported by the National Natural Science

Foundation of China (32070011).

784 ... Zhang & al.

Literature cited

Archer AW. 1993. A chemical and morphological arrangement of the lichen genus Pertusaria.

Bibliotheca Lichenologica 53: 1-17.

Archer AW. 1997. The lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 69: 5-249.

Archer AW, Elix JA. 2013. New species of Pertusaria (Pertusariaceae) from Australia and New

Zealand. Telopea 15: 111-117. https://doi.org/10.7751/telopea2013015

Archer AW, Elix JA. 2018. New combinations of Australian species in the genus Lepra Scop.

Australasian Lichenology 82: 130-136.

Elix JA. 2014. A catalogue of standardized chromatographic data and biosynthetic relationships for

lichen substances, 3rd ed. Published by the author, Canberra.

Galloway DJ. 2007. Flora of New Zealand: lichens, including lichen-forming and lichenicolous

fungi, vol. 2: Pannaria - *Zwackhiomyces. Manaaki Whenua Press, Lincoln, New Zealand.

Hafellner J, Tiirk R. 2016. Die lichenisierten Pilze Osterreichs — eine neue Checkliste der bisher

nachgewiesen Taxa mit Angaben zu Verbreitung und Substratoékologie. Stapfia 104. 216 p.

Lendemer JC, Harris JC. 2017. Nomenclatural changes for North American members of the

Variolaria-group necessitated by recognition of Lepra (Pertusariales). Bryologist 120: 183-190.

https://doi.org/10.1639/0007-2745-120.2.183

Ren Q. 2019. Taxonomic revision of the genus Lepra (Pertusariales) in China. Mycosystema 38(11):

1840-1864. https://doi.org/10.13346/j.mycosystema.190150

Ren Q, Sun ZS, Zhao ZT. 2009. Pertusaria wulingensis (Pertusariaceae), a new lichen from China.

Bryologist 112(2): 394-396. https://doi.org/10.1639/0007-2745-112.2.394

Schmitt I, Lumbsch HT. 2004. Molecular phylogeny of the Pertusariaceae supports secondary

chemistry as an important systematic character set in lichen-forming ascomycetes. Molecular

Phylogenetics and Evolution 33(1): 43-55. https://doi.org/10.1016/j.ympev.2004.04.014

MYCOTAXON

October-December 2021— Volume 136, pp. 785-788

https://doi.org/10.5248/136.785

Validation of five Peronospora species names

WILLIAM J. DAvIs*? & JO ANNE CROUCH"

' United States Department of Agriculture, Agricultural Research Service,

Mycology and Nematology Genetic Diversity and Biology Laboratory,

10300 Baltimore Avenue, Beltsville, Maryland 20705, U.S.A.

? Oak Ridge Institute for Science and Education,

ARS Research Participation Program, Oak Ridge, Tennessee, U.S.A.

*CORRESPONDENCE TO: joanne.crouch@usda.gov

ABSTRACT—Some Peronospora names included in the last major taxonomic treatment (a

1991 annotated list of names) were invalid because the author failed to provide the Latin

description required at the time of their proposal. Here five such names—Peronospora

coronillae-minimae, P. erini, P. papaveris-pilosi, PR. ranunculi-flabellati, and P. viciicola—are

validated either by providing an English description or by reference to the original English

description.

Key worps—biodiversity, downy mildew, Peronosporaceae, Peronosporales, plant pathogens

Introduction

Peronospora (Oomycota, Peronosporales, Peronosporaceae) is a species-

rich genus of biotrophic plant pathogens that cause downy mildew diseases

(Constantinescu 1991, Thines & Choi 2016, Salgado-Salazar & al. 2018). It

is also a nomenclaturally and taxonomically complicated genus (Shaw 1981,

Skalicky 1983, Constantinescu 1991). Broadly, there are two approaches to

describing species in Peronosporaceae. One approach applies a broad species

concept, with one species of Peronospora infecting many hosts within a

single plant family. The other approach applies a narrow species concept,

with species having narrow host specificity (Skalicky 1983, Constantinescu

1991). Molecular phylogenetic research seems to validate the application

of a narrow species concept (e.g., Thines & Choi 2016), even though the

786 ... Davis & Crouch

morphology of phylogenetically distinct species may overlap (e.g., Garcia-

Blazquez & al. 2008; Petrzelova & al. 2017).

Constantinescu (1991) provides the most comprehensive list of

Peronospora names and is an invaluable resource to those studying the

genus. In his list, Constantinescu (1991) included many names that were

invalidly published because the describing author failed to clearly designate

a holotype specimen. Examples include several Peronospora taxa described

by Gaponenko (1972), who did not designate individual type specimens,

that have been recently validated by Thines (2019). Other names are invalid

because the describing authors did not provide a description in Latin, which

was required by the nomenclatural code in effect at the time of publication

(Constantinescu 1991). The aim of this paper is to validate Peronospora

coronillae-minimae, P. erini, P. papaveris-pilosi, P. ranunculi-flabellati, and

P. viciicola, for which holotypes were designated but a Latin description

was not provided. Here an English description is provided as allowed by

the most recent International Code of Nomenclature for algae, fungi, and

plants (Turland & al. 2018).

Taxonomy

Peronospora coronillae-minimae Vienn.-Bourg. ex

WJ. Davis & J.A. Crouch, sp. nov.

MB 838139

“Peronospora coronillae-minimae” Vienn.-Bourg., Bull. Soc. Mycol.

France 69: 332. 1954 [nom. inval., ICN (Shenzhen) Art. 39.1]

CONIDIOPHORES in scattered tufts on abaxial side of Coronilla minima

leaves, emerging in clusters of 3-6, 160-200 um long with a cylindrical

or clavate trunk 8-10 um thick. STERIGMATA forming at an obtuse angle.

Conip1a subhyaline to pale yellow, ellipsoid, 19-32 x 16-28 um (average:

25 x 21 um). OosporEs numerous, pale yellow, smooth wall 2.5 um thick,

25-36 X 23-34 um.

Type: France, Seine-et-Oise, near Ferté-Allais, Coronilla minima L., May 1952, G.

Viennot-Bourgin, (Holotype, PC).

Peronospora erini Vienn.-Bourg. ex W.J. Davis & J.A. Crouch, sp. nov.

MB 838140

“Peronospora erini’ Vienn.-Bourg., Bull. Soc. Mycol. France 69:

334. 1954. [nom. inval., ICN (Shenzhen) Art. 39.1]

CONIDIOPHORES scattered, hydrophilic, on margin of yellowing leaves,

grayish, ragged, 80-130 um long, 6.9 um diam., branching in upper 1/2 to 1/3;

Validation of five Peronospora names ... 787

ultimate branchlets contracted. Conip1a grey, ellipsoid, slightly apical, 12-31

x 12-21 um (average: 24.2 x 17.5 um). OOsPpoREs unknown.

Type: France, Hautes-Pyrénées, Cirque de Gavarnie, small bog, Erinus alpinus L.,

August 1952, G. Viennot-Bourgin (Holotype, PC).

Peronospora papaveris-pilosi Vienn.-Bourg. ex W.J. Davis & J.A. Crouch, sp. nov.

MB 838141

“Peronospora papaveris-pilosi’ Vienn.-Bourg., Bull. Soc. Mycol. France

69: 335. 1954. [nom. inval., ICN (Shenzhen) Art. 39.1]

Forming angular, yellow-brown spots (confined by veins on Papaver pilosum

leaves), 4.2 mm diam. Mycelium homogenous, dense, brownish, on abaxial

side of the spots. CONIDIOPHORES erect, 300-400 um long, divided, branched

in upper half. Secondary branches bearing moderate to few divisions.

STERIGMATA forming at an acute angle. Conip1A ellipsoid or globose, pale

brown, 15-35 x 13-24 um (average: 25.5 x 21.4 um). OosPporEs unknown.

Type—France, Seine-et-Oise, Grignon Botanical Garden, Papaver pilosum Sm., April

1931, G. Viennot-Bourgin (Holotype, PC).

Peronospora ranunculi-flabellati Vienn.-Bourg. ex

WJ. Davis & J.A. Crouch, sp. nov.

MB 838142

“Peronospora ranunculi-flabellati” Vienn.-Bourg., Bull. Soc. Mycol.

France 69: 335. 1954. [nom. inval., ICN (Shenzhen) Art. 39.1]

Mycelium dense, grayish brown on abaxial side of Ranunculus paludosus

leaves. CONIDIOPHORES variable, 300-450 um long, branching slightly below

upper half. Secondary branches distinct, flexible, branching dichotomously

1-2 times. STERIGMATA short and acute. Conrp1a grayish brown, averaging

19 x 17.4 um. Oospores numerous, yellowish walls, slightly pleated,

20-25 um diam.

Type: France, Seine-et-Oise, Grignon Botanical Garden, Ranunculus chaerophyllos

[redet. as R. paludosus], May 1937, G. Viennot-Bourgin (Holotype, PC).

Peronospora viciicola L. Camp. ex WJ. Davis & J.A. Crouch, sp. nov.

MB 838143

“Peronospora viciicola” L. Camp. [as ‘vicicola’], Bull. Wash. State Agric.

Exp. Sta. 318: 18. 1935. [nom. inval., ICN (Shenzhen) Art. 39.1]

English description provided in Campbell (1935: 18).

Type: United States of America, Washington, Steamboat Rock, Vicia gigentea Hook.

[= V. nigricans subsp. gigantea (Hook.) Lassetter & C.R. Gunn], 7 September 1934, L.

Campbell (Holotype, WSP 16631).

788 ... Davis & Crouch

Acknowledgments

We appreciate Mme. Josette Rapilly, bibliothéque ala SME for her help in retrieving

a copy of Viennot-Bourgin (1954). We thank Amy Rossman (retired, USDA, USA)

and Nina Shishkoff (USDA, USA) for their insightful reviews. This research was

supported in part by an appointment of William J. Davis to the ARS Research

Participation Program administered by the Oak Ridge Institute for Science and

Education (ORISE) through an interagency agreement between the US Department

of Energy (DOE) and USDA. ORISE is managed by ORAU under DOE contract

number DE579AC05-06OR23100. Mention of trade names or commercial products

in this publication is solely for the purpose of providing specific information and does

not imply recommendation or endorsement by the US Department of Agriculture.

The USDA is an equal opportunity provider and employer.

Literature cited

Campbell L. 1935. Downy mildew of peas caused by Peronospora pisi (DeB.) Syd. Bulletin,

Washington State Agricultural Experiment Station 318.

Constantinescu O. 1991. An annotated list of Peronospora names. Thunbergia 15. 110 p.

Gaponenko NI. 1972. The family of Peronosporaceae of central Asia and south Kazakhstan: a

classification key (Translated from Russian). Fan Publishing House of the Uzbek SSR, Tashkent.

341 p.

Garcia-Blazquez G, Géker M, Voglmayr H, Martin MP, Telleria M, Oberwinkler FE. 2008. Phylogeny

of Peronospora, parasitic on Fabaceae, based on ITS sequences. Mycological Research 112:

502-512. https://doi.org/10.1016/j.mycres.2007.10.007

Petrzelova I, Choi Y-J, Jemelkova M, Dolezalova I, Kruse J, Thines M, Kitner M. 2017. Confirmation

of Peronospora agrimoniae as a distinct species. European Journal of Plant Pathology 147:

887-896. https://doi.org/10.1007/s10658-016- 1058-8

Salgado-Salazar C, Shishkoff N, Daughtrey M, Palmer CP, Crouch JA. 2018. Downy mildew:

a serious disease threat to rose health worldwide. Plant Disease 102: 1873-1882.

https://doi.org/10.1094/PDIS-12-17-1968-FE

Shaw CG. 1981. Taxonomy and evolution. 17-29, in: DM Spencer (ed.). The Downy Mildews.

Academic Press, London, UK.

Skalicky V. 1983. The revision of species of the genus Peronospora on host plants of the family

Rosaceae with respect to Central European species. Folia Geobotanica et Phytotaxonomica 18:

71-101. https://doi.org/10.1007/BF02855638

Thines M. 2019. Fixing loose ends in downy mildew research—the Peronosporaceae of Kazakhstan

by Nina Ivanovia Gaponenko. Schlechtendalia 36: 133-139.

Thines M, Choi Y-J. 2016. Evolution, diversity, and taxonomy of the Peronosporaceae, with focus

on the genus Peronospora. Phytopathology 106: 6-18. https://doi.org/10.1094/PHY TO-05-15-

0127-RVW

Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS & al. 2018.

International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted

by the Nineteenth International Botanical Congress, Shenzhen, China, July 2017. Regnum

Vegetabile 159. https://doi.org/10.12705/Code.2018.

Viennot-Bourgin G. 1954. Notes mycologiques (III). Bulletin de la Société Mycologique de France

69: 332-342.

MY COTAXON

October-December 2021—Volume 136, pp. 789-818

https://doi.org/10.5248/136.789

Typification of Agaricus cespitosus, Ag. oniscus,

and Ag. sphagnicola and their synonymy

with Lichenomphalia umbellifera

ANDRUS VOITK

13 Maple St., Humber Village, NL, Canada, A2ZH 2N2

*CORRESPONDENCE TO: seened@gmail.com

ABSTRACT—Protologue descriptions and original material of Agaricus cespitosus,

Ag. oniscus, and Ag. sphagnicola were studied to determine the intended species concept

for each. Agaricus cespitosus fits the second most common presentation of the current

Lichenomphalia umbellifera and is placed in synonymy with it. Agaricus oniscus was

created as a synonym of Ag. cespitosus, confirmed by its original material; therefore, it is

also placed in synonymy with L. umbellifera, resolving problems noted with past attempts

trying to apply the epithet to darker sphagnicolous species of Arrhenia. Careful analysis

of the protologue for Ag. sphagnicola revealed a good fit with L. umbellifera but serious

conflict if applied to species of sphagnicolous Arrhenia; it was, therefore, also synonymized

with L. umbellifera. Agaricus cespitosus and Ag. oniscus are lectotypified with illustrations

from their original material, Ag. oniscus is epitypified with a modern sequenced collection

from Sweden, and Ag. sphagnicola neotypified with a K collection made by Berkeley, both

identified as L. umbellifera.

Key worps—Omphalina ericetorum, nomenclature, taxonomy

Introduction

My first task would certainly be to rectify the names. ... If the names are not correct, if

they do not match realities, language has no object. If language is without an object, action

becomes impossible—and therefore, all human affairs disintegrate and their management

becomes pointless and impossible. Hence the very first task of a true statesman is to rectify

the names.

— THE ANALECTS OF CONFUCIUS, tr. Simon Leys (1997)

790 ... Voitk

In preparation for of a study of sphagnicolous species of Arrhenia Fr. (hereafter,

sphagnicolous arrhenias), epithets applied to this group were reviewed. This

revealed that some past workers have questioned the application of two

epithets, Agaricus oniscus and Ag. sphagnicola to this group, referring at least

the latter to the species currently known as Lichenomphalia umbellifera. This

study was undertaken to elucidate the original species concepts for these and

allied taxa.

Lichenomphalia umbellifera (L.) Redhead & al.

Over the years this species has been assigned to several genera and known

by several epithets. Part of its convoluted nomenclatural history has been

reviewed by Redhead & al. (2002), concluding with an argument for the current

name, which has resulted in welcome nomenclatural stability. Molecular

studies (Geml & al. 2012) have revealed that despite its varied appearance, in

the Northern Hemisphere L. umbellifera is a single species, not a complex. Its

mercurial nature notwithstanding, it is readily separated from sphagnicolous

arrhenias, although not all characters that separate them are well known.

In the Canadian province of Newfoundland and Labrador (NL) where I

live, L. umbellifera is ubiquitous. Between 2003 and 2016, I made 32 collections

of it throughout the province, and Foray Newfoundland & Labrador (FNL,

the provincial mushroom club) recorded it 73 times. Mt Ignoble, a barren

hilltop less than 1% hours on foot from our door, is the closest site where I

can find it with regularity. This small lichenized omphalinoid species has very

variable macroscopic appearance, its colour varying from near-white (Fic.

1A,C), ochraceous (Fic. 1B), yellowish with some purple on the stipe (Fic.

1E), and beige-tan (Fic. 1D); often the membranaceous pileus has centripetally

widening bands of grey in translucent areas (Fic. 1A,B). The shape of the

pileus varies from the less common convex with adnate gills (Fic. 1A,B,D,F),

to markedly upturned or infundibuliform (Fic. 1C,E), with upturned edges

and downsloping gills. This variation may lead one to suspect two different

species, but likely the shape is related to environmental factors and age. A

plane pileus with downturned edges and adnate gills is the probable result

of a pileus growing freely in concert with the hymenium. Hardening of the

pileipellis from exposure to sun and wind would impede its growth, and active

lamellar growth would then likely produce funnel-shaped pilei with upturned

(downsloping) gills. In all but the most protected moist conditions this process

would be repeated over time, making the funnel shape (seen in about 70% of

our collections) increasingly common as the basidiomes age. The habitat of

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 791

Fic. 1. A-F: Lichenomphalia umbellifera in NL. See text. E: Photo: Michael Burzynski.

L. umbellifera ranges from arctoalpine barrens (Fic. 1 A,D), bog (Fig. 1C), to

lowland forest (Fic. 1E). It grows on a variety of substrates, earth, Sphagnum

L. or moss, wood, with peat seemingly preferred. The species is known for

two classical presentations, one resembling soldiers in a row on moss-covered

rotting logs (Fic. 1E) and the other in great troops on the wet sides of peat pits

(Fic. 1F).

Agaricus cespitosus Bolton

James Bolton (1735-1799) was an English naturalist whose illustrated

multi-volume AN HISTORY OF FUNGUSSES GROWING ABOUT HALIFAX is the

first English book devoted to fungi. In volume 1 Bolton (1788) describes a

new species, Agaricus cespitosus. The description matches that of the current

L. umbellifera, particularly the yellowish clay colour and the iconic growth

in troops from the sides of peat pits. Before publishing his book, Bolton

produced a six-volume pictorial record of the fungi he had encountered, with

handwritten notes and aquarelles of the species. Agaricus cespitosus appears

in volumes 1 (Bolton 1784) and 3 (Bolton 1786) with a different aquarelle in

each (reproduced here as Fics 2A,B). The aquarelle from the third volume

792. ... Voitk

Siti DobiMiaede ora OAS

Fic. 2. Agaricus cespitosus: illustrations by Bolton. A: From vol 1 of his pre-publication

illustrations (Bolton 1784). B: From vol 3 of the pre-publication illustrations (Bolton 1786), here

declared lectotype for the species. C: The definitive illustration (Bolton 1788, tab. XLI, fig. C),

cited by Fries as an illustration of Ag. oniscus and selected here as the lectotype for that species.

Some volumes were hand coloured by Bolton; other volumes and editions were coloured by other

people and various techniques. This uncoloured image was selected to avoid potential problems

from differences in colouring. Images in the public domain; A & B, courtesy Special Collections,

USDA National Agricultural Library; C, courtesy The New York Botanical Gardens Library.

of his Icones (reproduced here as Fic. 2B) became the basis for the copper

engraving in his published work (Bolton 1888, tab. XLI, fig. C). That figure is

reproduced here as Fic. 2C. No type tissue remains, making these illustrations

the only available original material for lectotypification. All three illustrations

are accurate representations of L. umbellifera, particularly the umbrella shape

of radiating wedges and the representation of both convex and upturned pilei.

Testing my interpretation of Bolton’s formal illustration for Ag. cespitosus,

I sent the reproduced Fic. 2C to an expert panel of mycologists familiar

with arctic-alpine mycota: Jozsef Geml, Gro Gulden, Pierre-Arthur Moreau,

Esteri Ohenoja, and Anna Ronikier. Each mycologist was asked what species

the drawing represented, without providing a reason for the question, any

suggestion of region, habitat or other information, or any potential choice

of names. One panelist identified the drawing as Entoloma sericellum (Fr.)

P.Kumm. and one as Omphalina pyxidata (Bull.) Quél., while three immediately

concluded it was Lichenomphalia umbellifera. The first two were asked for

the best match out of three names, Arrhenia oniscus (Fr.) Redhead & al.,

Arr. philonotis (Lasch) Redhead &al., L. umbellifera; both selected L. umbellifera.

Agaricus cespitosus Bolton, Hist. Fung. Halifax 1: 41. 1788.

Type: Holotype: unavailable. Lectotype: (here designated, MBT10004616): Bolton,

Agaricus cespitosus, cones Fungorum circa Halifax sponte nascientum vol. 3, p. 103,

1784, in the book held in Special Collections, USDA National Agricultural Library;

[reproduced here as Fic. 28].

= Lichenomphalia umbellifera (L.) Redhead & al.

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 793

CoMMENT: Bolton’s second illustration (reproduced here as Fic. 2B) is chosen

as lectotype because it is the original image on which the protologue illustration

(reproduced here as Fic. 2C) is based. Because both protologue, lectotype and

other illustrations all fit accurately with L. umbellifera, Ag. cespitosus is placed

in synonymy with it, as a later synonym.

Agaricus oniscus Fr.

In his protologue of Ag. oniscus, Fries (1818) cited only the description and

illustration of Agaricus cespitosus as a previous description of the same species.

Fries declared that his name Ag. oniscus includes all other names (“et ind. univ.’)

and excludes—in the sense of replaces—all synonyms (“excl. omn. synon.”), i.e.,

even the earlier Ag. cespitosus of Bolton. By citing only Bolton's description, and

emphatically declaring Bolton’s name a synonym of Ag. oniscus, Fries leaves

no doubt about his certainty that they are conspecific. Ordinarily a later name

would be declared superfluous but Fries redescribed Ag. oniscus in his Systema

(Fries 1821), whereby the name became sanctioned, giving it precedence over

Bolton’s earlier Ag. cespitosus. Thus, application of the epithet oniscus should

be very straightforward: in his protologue Fries stated that his species is the

same as Bolton's, but because the Fries epithet became sanctioned, the species is

now known as Ag. oniscus. Because Bolton's species is a later description of the

current L. umbellifera, epitypified with a sequenced specimen of that species

(vide supra), Ag. oniscus is also a later synonym of the current L. umbellifera.

However, confusion was instilled into this seemingly straightforward

relationship by Fries himself, who in the course of the subsequent half century

markedly altered his concept of the species to which he applied the epithet

oniscus. Already three years after the 1818 protologue Fries (1821) said that since

its introduction he had come to view Ag. oniscus as perhaps (“forsan”) a variety

of Ag. cespitosus. This is the first of progressively greater deviations that ended

with Fries’s application of the epithet to a taxon markedly different from the

one described in the protologue. Twenty years after the protologue Fries (1838)

described its originally non-decurrent gills as adnate to decurrent (“adnato-

decurrentibus”), and 36 years after the protologue Fries (1854) stated that this

erstwhile synonym for Bolton’s turf agaric was rare in mossy and turfy habitats

but common beside the sandy paths of the botanical gardens in Slottsbacken

near Uppsala. Thirteen years after that Fries (1867) applied oniscus to a dark

(“obscure”) species with a convex-umbilicate to infundibuliform (“convexo-

umbilicatus |. infundibuliformis”) pileus, decurrent ash grey (“cinereus”

lamellae, and grey (“griseus”) stipe. His evolving concept had veered so much

794 ... Voitk

from his early description and Bolton's species that by 1867 he was moved to

offer a more characteristic image (“iconem formae typicae”) of his new concept

(reproduced here as Fic. 3A).

Jules Favre, the pioneer of alpine fungal ecology (Brunner & al. 2017), took

this a step further, stating that Fries’s 1867 illustration was too light (Favre 1948).

At the time the rules governing nomenclature declared Fries’s Systema of 1821

as the starting point for agaric names, denying earlier names nomenclatural

priority. Favre complied with these rules and used the name oniscus, treated

by Fries in 1821, not cespitosus or some earlier name cited by him. Why, then,

did he not use the description of Ag. oniscus from the protologue, essentially

duplicated in the Systema, rather opting for Fries’ description of 1867, a

description at odds with the original material, and clearly of a different species?

We may never know the answer, but two potential contributing factors have

been described to me. Pierre-Arthur Moreau (private communication) related

that in 1948, French authors considered Fries’s 1821 Systema a youthful work,

and looked on the 1867 Monographia as the authoritative opus—so much so,

that at one time French mycologists proposed that the Monographia, rather

than the Systema, should be the sanctioning work for Hymenomycetes. Favre's

friend and colleague, Marcel Josserand, who had great influence on Favre's tenets

in these matters, was of this view. Henning Knudsen (private communication)

pointed out that at the time there were very few copies of the Systema available

in Europe, whereas the Monographia was common. Generally Fries remained

true to his original descriptions in subsequent treatments, so that most of the

time the description of a species in the Monographia did not differ significantly

from the one in the Systema. Therefore, as a rule the later description could be

used confidently, should the earlier one not be at hand. Whether these insights,

alone or in combination, played a role in Favre's treatment of Ag. oniscus is

a matter of speculation, but Favre did leave on record an illustration of his

collection in G, including three near-black specimens (reproduced here as

Fic. 3B), to depict his concept of the species.

Favre's study of Jurassic wetlands, where he treated Ag. oniscus, was followed

by intense exploration of the alpine zone, giving him an unparalleled mastery

of these mycotas. The resultant regard for his opinions helped usher in an era

of applying Ag. oniscus to really dark species in Europe. His influence persisted

until distance and time permitted Bigelow (1985), in North America 37 years

later, to comment on the “discordant” fit of Favre's interpretation with Fries’s

protologue, stating, “Favre may not have had the same species as Fries.” Indeed,

rather than accepting Favre's interpretation of Ag. oniscus, Bigelow (1958)

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 795

Fic. 3. Brown arrhenias, arenicolous (A), and sphagnicolous (B-G), scaly-capped (D-G, possibly

C), smooth-capped (A, B). A: Fries’ 1867 image of Agaricus oniscus, at that time considered by him

to be a non-sphagnicolous arenicolous species, darker and evenly brown, significantly different

from Bolton’s image that Fries cited as an unquestioned synonym in 1821. Image in the public

domain. B: Jeanne Favre's illustrations for Jules Favre's collections of Omphalia oniscus in G. Image

courtesy of the Conservatoire et Jardin botaniques de Geneve. C: Quélet’s collection, “Agaricus

(Omphalia) oniscus Fr” from the Juras Mountains in the E. Fries Herbarium of UPS (F-012426);

seemingly sphagnicolous, but fits with a species of Arrhenia and not with Bolton's Ag. cespitosus.

Photo courtesy of UPS. D: Photo of a scaly-capped sphagnicolous arrhenias in NL. E: Peck’s

aquarelle of the surviving of two syntypes of Agaricus gerardianus (NYS-1339-d). Image (I-1390)

courtesy NYS. F: Jakob Lange's illustration of Omphalina sphagnicola. Image courtesy of Henning

Knudsen, permission courtesy of Lene Lange. G: Jeanne Favre's illustrations for Jules Favre's

description of Omphalia sphagnicola. Image in the public domain. NOTE: some of the elements

on these illustrations were moved to fit space requirements, and sharpness was augmented to

exaggerate scale detail beyond the original.

796 ... Voitk

elected to apply Persoon’s epithet icmadophilus (Persoon 1801) to what he

thought was Fries’s original Ag. oniscus, elevating Agaricus epichysium var.

icmadophilus Pers. to species as Clitocybe icmadophila (Pers.) H.E. Bigelow.

Redhead (1979) followed Bigelow in the use of this epithet, transferring it to

Omphalina as O. icmadophila (Pers.) Redhead. Despite his justified misgivings

about Favre's interpretation, 27 years later Bigelow (1985) relinquished the

epithet icmadophilus, because he considered its protologue description “very

short and could also be of several species” and justified accepting Favre's dark

concept because of Fries’s later volte-face to a much-altered concept of Ag.

oniscus. Left alone using icmadophilus, Redhead did not make public a change

of mind, but stated (personal communication) that he “acquiesced to standard

usage” by 2002 (Redhead & al. 2002). This was the year Ag. oniscus—but not

O. icmadophila—was transferred to Arrhenia’.

Since Favre's time, rules governing nomenclature set forth in the International

Code of Nomenclature for algae, fungi, and plants (henceforth the Code;

Turland & al. 2018), have evolved. Before turning to the Code to reconcile

conflicting opinions of which name to apply to a species first described over

230 years ago, some due diligence seems prudent to ensure that this does not

produce unsuspected conflict. The following paragraphs explore whether the

descriptions of Ag. cespitosus and Ag. oniscus fit with each other, whether both

fit the concept of the current L. umbellifera, whether discordant elements in the

descriptions can be reconciled with L. umbellifera, whether the introduction

of a new name or different rankings are compatible with the synonymization,

whether these descriptions fit better with sphagnicolous arrhenias, whether

synonymy causes significant instability of established practice, and, finally,

determine all type material available.

The descriptions of Bolton (some elements taken from his illustration) and

Fries are very similar (TABLE 1). Bolton describes his species from the sides

of peat pits, and Fries states his comes from montane heaths. The differing

habitats are not mutually exclusive but indicate that Fries is not making a

mechanical name change and is personally familiar with the species. He states

as much in the Systema (“v. v.’), where he provides mossy sites and old logs

(“truncos venustos”) as an additional substrate (classical L. umbellifera habitat/

‘Although sometimes rendered “Arrhenia/Omphalina onisca’, the somewhat counterintuitive-

sounding Arrhenia/Omphalina oniscus is correct. Fries chose the noun Oniscus, the genus to which

the woodlouse belongs; there is no corresponding Latin adjective. To make certain his intent was

clear, he capitalized the epithet, as was the custom of the time for proper nouns used as epithets

(e.g. generic names). A noun used as an epithet retains its gender, regardless of the gender of the

genus name that it modifies.

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 797

TABLE 1. Protologue characters of Agaricus cespitosus and Ag. oniscus

BOLTON (1788) Agaricus cespitosus Fries (1818) Agaricus oniscus

COLOUR Yellowish clay Dirty or greyish

GROWTH Gregarious Gregarious

HABITAT Sides of peat pits Montane heaths

PILEUS Striate, translucent, smooth, 25 mm diam., Striate, translucent, smooth, 25 mm

convex in youth, then plane, and diam., convex in youth, then plane,

eventually upturned; not umbilicate and eventually depressed; not

unless totally upturned described as umbilicate

LAMELLAE Distant, adnate, not decurrent (excepting Distant, adnate (“adnatis”), not decurrent

the downsloping of upturned pilei). (specifically stipulated three years

later)

STIPE 25 mm high, cylindrical, usually bent to 25 mm high, cylindrical, often bent, at

wavy times wavy

substrate). The only difference worthy of comment is an apparently divergent

description of colour. Bolton’s is “yellow clay,’ while Fries used “lividus,” usually

a lighter leaden or bluish grey, which Bigelow (1985) found “vague and difficult

to visualize and reconcile”. Bolton's first prepublication aquarelle (reproduced

here as Fic. 4A) may provide some insight into the greys as Fries saw them.

The pileus has opaque radiating lines over the lamellae and lamellulae and

intervening lighter grey spaces over the thinner membranous parts. Similar

narrow translucent grey wedges can be seen in membranaceous species, like

L. umbellifera Fic. 4B, widening with age and increased hydration (Fic. 4C);

both opaque and translucent parts have a greyish cast. Both Fries and Bolton

saw grey in these specimens, and Bolton's description “yellow clay” was available

to Fries when he declared the two names synonymous, so that the use of either

colour should not be a major conflict.

Bolton’s English name, peat agaric, gives away the meaning of his Latin

epithet: ceespes is Latin for peat, and cespitosus is the adjective peaty. This

certainly fits with L. umbellifera, for which peat is a favourite substrate. Although

renaming organisms already described and named was not unusual at the

time, it seems likely that Fries renamed this species in the interest of clarity.

In mycology, commonly the adjective cespitosus has been used to indicate a

species with a several stipes arising from one “root”. In his protologue, where he

stated that oniscus is meant to replace all previous names, Fries did not say why

he replaced the name, but emphasized that the species is gregarious yet never

cespitose (“gregarius numquam cespitosus”) by making this the first sentence

of the description. It is not part of the section dealing with habitat, indicating

798 ... Voitk

that to Fries the adjective refers to morphology, not substrate. Having made this

precision in the protologue, Fries never described Ag. oniscus as non-cespitose

again in subsequent treatments, suggesting that this distinction was used solely

to explain his reason for renaming the species. Fries spoke Latin with his father

at home since childhood, wrote almost all of his books (including a layman's

field guide to mushrooms!) in Latin, and carried out the vast majority of his

foreign correspondence in Latin, so that he knew the language well. It may be

that he sought to avoid confusion, more than to “correct” Bolton’s Latin. In

either case, it is reasonable to assume that he took care to select a characteristic

and unambiguous new epithet.

Wood lice are common crustaceans in Sweden. Of the two genera, Oniscus

L. and Armadillidium Brandt, the former are much more common, coloured

various shades of grey, and do not roll up (Fic. 4D). Armadillidium species (pill

bugs) are dark to blackish, and roll into a wheel when in danger, so that from

the side the overlapping exoskeleton segments resemble the radially triangular

segments of the pileus (Fic. 4E). Armadillidium was split from Oniscus in

1833, so that at the time of Fries’s description both were considered Oniscus.

Hence, the name could suggest that both the centripetally radiating triangular

segments of the pill bug and the grey of the wood louse resemble the radial

segments and their grey translucent areas on the pileus of L. umbellifera.

Fries (1821) placed Ag. umbelliferus L. in Mycenaria (with decurrent

gills), and Ag. oniscus in Collybaria (with adnate gills) in his Systema. If

the species treated by Fries as Ag. umbelliferus is the same as the current

L. umbellifera—a subject beyond the scope of the current dissertation—then

its ranking should be no impediment to accepting their synonymy. While

we accord Fries’s sanctioned names priority over earlier ones, we do not do

the same for his taxonomy—even though he gave us our taxonomic system

of ranking. Taxonomy changes constantly in response to new information,

making it perfectly acceptable, for example, to place taxa in synonymy that

were originally separated by gill arrangement, should new information show

this to be an intraspecific, not interspecific, character. Such an outcome should

not be unexpected with a synonym of the current L. umbellifera, which has

basidiomata with both adnate and decurrent gills. Because taxonomy must

remain flexible, it is an inappropriate tool for nomenclature, which aims to

fix names so that they remain stable, regardless of any subsequent changes in

ranking.

Fries’s protologue fits well with the current L. umbellifera, an iconic species

in montane heaths, classically fruiting in troops on old logs (Fic. 1E) in

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 799

Fic. 4. Relationship of “oniscus” to pileal colour and form. A: Agaricus cespitosus (from Bolton's

lectotype colour illustration). B, C: Lichenomphalia umbellifera (C: older specimen). D: Oniscus

asellus L. in NL. E: Armadillidium vulgare Latreille (rolled up pill bug); Photo: Benjamin T.,

Wikimedia Commons, https://commons.wikimedia.org/w/index.php?curid=1936102 [Creative

Commons licence CC BY-SA 3.0].

woodlands, and on the sides of peat pits (Frc. 1F). Neither Bolton's nor Fries’s

description fits sphagnicolous arrhenias, which grow on living Sphagnum, not

peat, do not grow on wood (at least those species found in montane heaths),

make groups of 1-6, seldom troops, very quickly develop a pronounced deep

navel often without turning up the pileus, have deeply decurrent gills, and are

brown or grey-brown, not whitish yellow with light grey translucent zones.

Fries provided two different descriptions for his Ag. oniscus. In 1818,

he described a light, sphagnicolous species that does not fit a species of

Arrhenia, and by 1867 the description had changed to a dark arenicolous, non-

sphagnicolous Arrhenia. Applying Fries’ epithet to a third species that does

not fit with either of his two descriptions, namely a sphagnicolous species of

Arrhenia, produces irreconcilable conflict with both descriptions. It seems

much more reasonable to follow the Code, created to solve such matters, and

a) effect a solution that avoids conflict, b) is supported by the protologue, the

sanctioning treatment, and the lectotype, and c) respects Fries’s uncommon

degree of certainty about the synonymy of Ag. oniscus with Ag. cespitosus.

800 ... Voitk

A small number of names had been applied to an even smaller number of

sphagnicolous arrhenias with no apparent consistency until the work of Favre

(1948). After that, most authorities, e.g., Bigelow (1985) or Redhead (Redhead

& al. 2002), whether willingly or reluctantly, began to apply the name Arrhenia/

Omphalina oniscus, to a dark species. Reported spore sizes for Arr./O. oniscus

by selected students of these taxa (Favre 1948, Cleémencon 1982, Bigelow 1985,

Breitenbach & Kranzlin 1991, Kuyper 1995, Gminder 2001, Gulden 2005,

Elborne 2008) fall within a small range, 6-10 x 3.5-6 um in size (Fic. 5). This,

as well as the shared omphalinoid habitus, dark colour, and habitat suggest

that these authors probably apply the epithet to the same species. However,

other leading students of these taxa have identified similar species with

similar spore measurements using different epithets, such as Bigelow (1985):

Clitocybe icmadophila; Kuyper (1995): Omphalina philonotis (Lasch) Quél.;

Lange & Lange (1982): Omphalina epichysium (Pers.) Quel. In addition, Orton

(1960) described a dark species, Omphalina fusconigra Orton, with spore

measurements well outside this range.

Indeed, descriptions of the species whose spore measurements are shown

in Fic. 5 differ by more than their spore measurements. For example, Gulden

described the dark O. oniscus (Fr.) Quél. as an obligate sphagnophile, whereas

Favre described it as a facultative sphagnophile, a significant difference in a

group with possibly narrow photobiont association. The aquarelles painted by

his wife Jeannne to illustrate his collections suggest that Favre may have applied

the epithet to more than one species. His collections of O. oniscus deposited

in G are illustrated by opaque, non-striate smooth-capped basidiomes of two

widely different colours, one a yellowish ochre-tan, and the other a bluish

black-gray (reproduced here as Fic. 3B). The larger upper two basidiomes,

one of each colour, also illustrate his interpretation of the species (Favre 1948).

While different editions differ somewhat in the shading and it is possible that

the light colour is due to drying, near-black and yellowish tan are an unlikely

intraspecific colour variation. Finally, Arr./O. oniscus is the most common

name for sphagnicolous arrhenias in European herbaria (Robert Liicking,

personal communication). This is disproportionately high for the prevalence of

the species, again suggesting that the name is applied to multiple taxa.

Preliminary data from our ongoing investigation of sphagnicolous arrhenias

seem to support this view: to date we have identified only one really dark

species of sphagnicolous Arrhenia (an obligate sphagnophile, which I know

well because it is relatively common in the northern part of NL) whose spore

measurements (Fic. 5) match those reported by Elborne (2008). Presumably

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 801

Gulden 2005 Kuyper 1995

Bigelow 1985 a, Oo

. o

q 0)

5 S

afc es)

3 Favre 1948

Breitenbach & Kranzlin 1991 E|borne 2008

6 7 8 9 10

Fic. 5. Various major authors’ spore measurements of Arrhenia/Omphalina oniscus in um.

Length on x-axis and width on y-axis.

this is the species to which the name Arr. oniscus is most commonly applied

but, as mentioned, applying oniscus to facultative sphagnophiles at the same

time draws other species into the concept. Thus far, our preliminary studies

have found type collections of two different species in the phylogenetic clade of

dark obligate sphagnophiles, so that nomenclatural consistency is lacking even

here. Clearly, traditional usage is not unanimous, and any resolution will cause

temporary instability to a number of collections to which the epithet has been

applied. Hence synonymizing Ag. oniscus with the current L. umbellifera is less

likely to cause more instability than conserving the former.

Typification as we know it today was unknown at the time of Fries, and

preservation of type specimens uncommon. The only specimen in the E. Fries

Herbarium of UPS identified as Ag. oniscus is an undated collection (F-012426)

from the Juras Mountains sent by Quélet (Fic. 3C) and labelled, “Agaricus

(Omphalia) oniscus Fr.’ Because Quélet was born 14 years after the protologue

of Ag. oniscus was published, clearly this is not part of the original material.

Similarly, the only illustration of Ag. oniscus left by Fries (1867; reproduced

here as Fic. 3A) was published 49 years after the protologue, thus also not part

802 ... Voitk

of the original material—in fact, markedly at odds with it. In his protologue for

Ag. oniscus, Fries (1818) cited Bolton’s copper engraving of the synonymous

(as avowed by him) Ag. cespitosus (reproduced here as Fic. 2C). In his Systema

Fries (1821) cites his 1818 protologue and Bolton's copper engraving separately.

Therefore, for Fries (author of the name Ag. Oniscus), Bolton's illustration

remains the sole de facto type material for the taxon. As we learned in the

discussion of Ag. cespitosus, above, that image is an accurate fit for the current

L. umbellifera (confirmed by a panel of mycologists familiar with arctic-alpine

mycota) and suitable for formal lectotypification.

Agaricus oniscus Fr., Observ. Mycol. 2: 209. 1818.

Type: Holotype: unavailable. Lectotype (here designated, MBT10004612): Bolton,

illustration in An History of Fungusses Growing about Halifax, plate XLI (41), figure

C, 1788, in the book held by the library of the New York Botanical Gardens. Epitype

(here designated, MBT 10004614): Sweden, Jamtland, Undersaker, Trillevallen, Valliste,

63.2672°N 13.1515°E, at treeline on east slope of mountain, growing on disturbed soil.

5 July 2000, leg. Johan Rova 2501 (GB 0184058).

= Agaricus cespitosus Bolton

= Lichenomphalia umbellifera (L.) Redhead & al.

ComMENT: Because the lectotypes for Ag. oniscus and Ag. cespitosus depict the

same gathering, the names are unavoidable synonyms and the taxa conspecific.

Both have separately been found to represent L. umbellifera, making both later

synonyms of that name. To stabilize these names, Ag. oniscus is epitypified with

a modern sequenced Swedish specimen identified as L. umbellifera.

Agaricus sphagnicola Berk.

Scaly-capped sphagnicolous arrhenias (Fic. 3D) are common, yet the

first description and illustration of an indisputably scaly one (Peck 1873) is

Ag. gerardianus Peck (reproduced here as Fic. 3E). Almost 40 years earlier,

Miles Joseph Berkeley, (1803-1889), a vicar and cryptogamist credited

with laying the foundation to English plant pathology, described Agaricus

sphagnicola (Berkeley 1836) as a “pale funnel-shaped agaric” with a “minutely

squamulose” pileus, fruiting in Sphagnum. Although Berkeley’s was not a

description of an unequivocally scaly-capped sphagnicolous Arrhenia, some

workers took Berkeley’s word “squamulose” out of context to justify applying

sphagnicola to such species. Lange (1937) illustrated his interpretation of

Omphalia sphagnicola (Berk.) P. Karst. with an infundibuliform, somewhat

dark and long-spored sphagnicolous species with obvious umbilical scales

(reproduced here as Fic. 3F). Because Berkeley observed that “the whole plant”

was “tough and elastic’, and Lange specified that the species he described was

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 803

neither tough nor cartilaginous, clearly they did not apply the epithet to the

same species. Indeed, Lange’s species (reproduced here as Fic. 3F) is close to

Peck’s (reproduced here as Fic. 3E), matching scaly-capped sphagnicolous

arrhenias (Fig. 3D), and not the species illustrated by Bolton (reproduced here

as Fic. 2A-C). Twenty-two years later Favre (1948) followed Lange's concept,

and also applied it to scaly species (reproduced here as Fic. 3G), thereby

endorsing Ag. sphagnicola as an earlier name for a scaly-capped sphagnicolous

Arrhenia. Some students of these species favoured one name, some the other,

some used other names (e.g., O. philonotis, as Lange had done earlier) and some

entertained two scaly-capped sphagnicolous species.

This state of affairs came to an end when Redhead (1979) reported that

Berkeley's putative type collection of Ag. sphagnicola at K was Gerronema

ericetorum (Pers.) Singer [= L. umbellifera]|. He noted that the “dirty

ochraceous” colour and “minutely squamulose” pileus and stipe in Berkeley's

protologue have “... led many authors to believe that A. sphagnicola is an earlier

name for Omphalina gerardiana (q.v.); but concluded that the colour was well

within the range of the “variable” G. ericetorum and attributed the reported

squamules to the “deteriorated condition” of the “over mature” basidiomes.

Thus, after a critical review of the protologue, Redhead was able to explain away

apparent inconsistencies, and satisfy himself that Berkeley’s description fit the

current L. umbellifera. He synonymized Ag. sphagnicola with G. ericetorum

[= L. umbellifera], and applied Peck’s Ag. gerardiana (reproduced here as

Fic. 3E) to the scaly-capped sphagnicolous arrhenias (Fic. 3D). Eight years

later, with the help of Thomas Kuyper (Redhead & Kuyper 1987), Redhead

discovered that the collecting label for Berkeley's putative type collection of

Ag. sphagnicola at K did not match the date and place of the type, as noted in

the protologue, and therefore, “... there is no guarantee that it represents the

original A. sphagnicola based on a specimen collected June 21, 1827, Chartley

Moss, Staffs, on Sphagnum acutifolium (Berkeley 1836).’ This discovery led

Redhead & Kuyper to state: “Currently Berkeley's species is recognized as a

distinct Omphalina, Omphalina sphagnicola (Berk.) Moser, in Europe. Based

on the uncertainty of the status of the specimen sent as type, the current and

traditional application of the name O. sphagnicola is maintained.”

At this time, the only published review of Berkeley’s protologue (Redhead

1979) stated that it fit with G. ericetorum [= L. umbellifera]. In their report

Redhead & Kuyper made no effort to re-examine the protologue, or withdraw,

contradict, or amend Redhead’s earlier conclusion, leaving standing the

position that the protologue was a better fit for L. umbellifera than for scaly-

804 ... Voitk

capped sphagnicolous arrhenias. Eventually Kuyper (1995) took up this

question, critically reexamined Berkeley’s protologue of Ag. Sphagnicola to

conclude that it “seems to fit better for Phytoconis ericetorum” [= L. umbelliferal]

than O. gerardiana (Peck) Singer. He found no reason to alter Redhead’s 1979

conclusions, rejected their joint intervening proposition to apply Ag. sphagnicola

to a scaly-capped sphagnicolous species, resynonymized Ag. sphagnicola

with the current L. umbellifera, and reapplied O. gerardiana to scaly-capped

sphagnicolous arrhenias. This would have settled the issue, had Redhead not

continued to apply Ag. sphagnicola to scaly-capped sphagnicolous arrhenias

(Redhead & al. 2002). With the erstwhile collaborators espousing different

directions, confusion reigned. Some, like I, followed Redhead’s opinion, and

continued applying Ag. sphagnicola to scaly-capped sphagnicolous arrhenias,

but the lack of a type specimen, identification of at least one Berkeley collection

of his Ag. sphagnicola as the current L. umbellifera, and Kuyper’s analytical

conclusions all added a growing number of converts to the cadre now applying

Arr. gerardiana (Peck) Elborne to these species.

Discussions with Scott Redhead had led me to follow his practice of treating

Arr. sphagnicola as a single scaly-capped sphagnicolous species, until our study

of bog arrhenias caused me to review the names and protologues of this group

and discover a vexing quagmire of discontent. Without type material, the

protologue, now the only source of a species concept, assumes even greater

importance in guiding the selection of new type material to fix the name.

Kuyper’s interpretation could only be tested by repeated analysis of Berkeley’s

description. While Kuyper had already done this, he published his conclusion

only and not the methodology or step-by-step analysis so that the reader could

judge the validity of his conclusions. I decided to suspend my own belief that

Ag. sphagnicola represents a scaly-capped sphagnicolous Arrhenia and test

Kuyper’s interpretation by analyzing each of Berkeley's key characters, one by

one. In NL, Lichenomphalia umbellifera is relatively common, and because I

am fond of its preferred habitats, I have encountered it often. Over 20 seasons

I had accumulated many collections from various areas of the province, with

31 photos of 21 collections. My plan was to review these 31 photos for the

presence or absence of each key character in Berkeley's protologue.

The strength of this approach is objectivity: these photos were taken at a

time when I used Arr. sphagnicola for scaly-capped sphagnicolous arrhenias

and was unaware of any controversy suggesting Arr. sphagnicola might be

conspecific with L. umbellifera. Photos were not taken to prove a point or

demonstrate the presence or absence of any specific characters, but merely

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 805

to document a collection. The weakness of this approach is that a photo

that documents a collection in a general way may not always reveal specific

distinguishing characters, and critical parts may be omitted or out of focus.

Thus, even with 31 photos to choose from, the odds are skewed slightly against

identifying characters of Ag. sphagnicola on these photos; finding Berkeley's

key characters under these conditions should constitute objective proof of

their presence in L. umbellifera. Discussion of each key character described by

Berkeley follows.

BASIDIOME: TOUGH AND ELASTIC. This character is not visible on photos,

of course, but is so significant it must be included: with this feature Berkeley

immediately eliminates the friable scaly-capped sphagnicolous arrhenias as

possible candidates for his epithet sphagnicola. For this reason Bigelow (1958)

stated that C. gerardiana, which he described as “always soft and fragile’,

“cannot be considered identical with Ag. sphagnicola Berk” Redhead (1979)

concurred, saying Bigelow “quite rightly pointed out” this incompatibility.

This character fits only with L. umbellifera and encounters a major and

irreconcilable conflict if applied to scaly-capped sphagnicolous arrhenias.

Just as with Lange's interpretation, the discussion is really over before we even

begin to look at images. The method I chose serves only to examine how well

other macroscopic characters support this conclusion.

PILEUS: DIRTY OCHRACEOUS. Berkeley’s English name for his species was

“pale funnel-shaped agaric’. In common usage, “light” indicates less intense

or diluted pigmentation (e.g., light blue), whereas “pale” indicates paucity or

lack of pigmentation (e.g., to become pale). Lichenomphalia umbellifera does

not have much colour, ranging from white through (dirty) yellow to nearly

mid-tan on occasion (Fics 1, 6). The Oxford English Dictionary renders

ochraceous as a pale hue. Both pale and dirty ochraceous fit the dingy

yellowish L. umbellifera. In contrast, scaly sphagnicolous arrhenias are brown

and patterned, with alternating radial stripes of light or dark brown. Both pale

and dirty ochraceous would be inexact terms to describe them. The colour fits

much better with L. umbellifera than sphagnicolous arrhenias.

PILEUS: BECOMING DARKER WITH AGE. Most of the darkening of

L. umbellifera occurs at the pileus rim and lamellar edges (Fic. 6B-D, M), which

show through the pileus due to its lighter hue and translucency. Sphagnicolous

arrhenias darken in a similar manner, and very occasional specimens undergo

an uncommon darkening reaction, not unlike that observed in the two smaller

specimens on Fic. 6A. Becoming darker with age fits with both L. umbellifera

and sphagnicolous arrhenias.

806 ... Voitk

PILEUS: INFUNDIBULIFORM. Variations of the infundibuliform shape

(Fic. 6A-J, L), with upturned pileus, deep navel, and downsloping gills, are

the most common presentation for L. umbellifera, seen in at least 70% of

NL basidiomes. Scaly-capped sphagnicolous arrhenias develop navels and

decurrent gills, also assuming a funnel shape, although usually with a broader

angle. “Infundibuliform” fits both L. umbellifera and an Arrhenia species.

PILEUS: FUNNEL-SHAPED FROM A VERY EARLY STAGE OF GROWTH. Early

disproportionate hymenial development relative to pileus growth gives the

cap-gill structure of immature L. umbellifera a triangular shape with an early

navel, leading to the funnel-shape (Fic. 6E-I). Hymenial growth occurs later

for arrhenias, which usually emerge as small discs with inturned edges and

hypoplastic gills, becoming umbilicate later in development. Early funnel

shape fits much better with L. umbellifera than sphagnicolous arrhenias.

PILEUS: FAINTLY STRIATE. Because all tissues are a light colour with little

contrast, striations of L. umbellifera are faint (Fic. 6A, H-L). Sphagnicolous

arrhenias with their incrusted pigment usually have markedly striate pilei.

Faintly striate fits much better with L. umbellifera than sphagnicolous

arrhenias.

LAMELLAE: THICK, WITH A FLAT EDGE, DIRTY OCHRACEOUS. Lamellae of

both L. umbellifera (Fic. 6A-D, L, M) and sphagnicolous arrhenias are thick,

with flat edges, but only those of the former are ochraceous. Thick, ochraceous

lamellae fit much better with L. umbellifera than sphagnicolous arrhenias.

STIPE: ON SPHAGNUM, ADHERING BY ITS DOWNY BASE. When L. umbellifera

grows in Sphagnum, its stipes usually seem instititious because they are

enveloped in Sphagnum (Fic. 6A-C), and at times a white mat of mycelium is

evident (Fic. 6A,B,M) but if teased out, a more prominently downy white base

can be seen (Fic. 6A,D-F). Sphagnicolous arrhenias also adhere to Sphagnum

but have a more easily seen downy base. Adherence to Sphagnum and a downy

base is present for both but may need seeking out with L. umbellifera more

than with sphagnicolous arrhenias.

STIPE: NEARLY IMPERVIOUS, HOLLOw. The protocol could not be followed

to examine the integrity of stipe context on my photos, because I had never

cloven the basidiomes to photograph stipe core. The basidiomes are so

small that sagittal bisection is inconvenient, and L. umbellifera is so readily

recognizable macroscopically that I had no reason to seek an additional

character for identification, particularly one of questionable value, given

that so many good field mycologists had already documented it as either

partly (subfistulosus; Fries 1815) or totally (Cooke 1883, Peck 1893, Bigelow

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 807

Fic. 6. Lichenomphalia umbellifera: characters of basidioma, pileus and lamellae. Dirty, ochraceous

pileus: A, G, I K, L. Darkening with age: B, C, D, M. Pileus infundibuliform: A-J, L. Pileus funnel-

shaped early: E-I. Pileus faintly striate: A, H-L. Lamellae thick with flat edge, dirty, ochraceous:

A-D, L, M. Pileus minutely squamulose: J-M.

1970, Breitenbach & Kranzlin 1991) hollow. Except for one collection that

I could not locate, the package containing all my L. umbellifera collections

sent for sequencing was lost in the mail on return. Thus, I had no voucher

tissue to examine. However, I had just returned from a collecting trip to Great

Caribou Island in Labrador (Voitk & Burzynski 2018), where L. umbellifera

is copious. Although participants had been asked to focus on other groups

and not “waste time” on L. umbellifera, six collections of the species had been

made. Of these six dried collections, one had stuffed stipes, one had partly

hollow ones, and four had obviously hollow stipes (Fic. 7G—K), one of them

impervious (Fic. 7G), and the other nearly so (Fic. 7H). Just in case the

drying process converts stuffed stipes to hollow, I decided to examine my next

808 ... Voitk

finds in situ. The very first fresh specimen I found after that also had a hollow

stipe (Fic. 71). All these specimens were mature, making it possible that stipes

of younger specimens may not be hollow. However, this is moot: finding six

out of seven consecutive random collections with hollow stipes, thereby

supporting similar observations by several respected mycologists (vide

supra), confirms that a hollow stipe fits with L. umbellifera and, therefore, is

not a useful differentiator for these taxa.

STIPE: VERY MINUTELY SQUAMULOSE. This is the key to unravelling

Berkeley's meaning of the scaliness of his taxon, and therefore deserves

detailed attention. Redhead (1979) quite correctly observed, “The presence of

squamules on both the stipe and pileus is certainly unusual for any Omphaloid

species from bogs.” In 20 years of surveying bogs and examining several

hundred basidiomes of L. umbellifera and sphagnicolous arrhenias, I have yet

to see one of either with a scaly stipe. Redhead’s (1979) explanation that such

scaliness might be due to an “over mature” specimen, whose “deteriorated

condition may explain the reported squamules” contradicts Berkeley's

description of the stipe as “... smooth, except at first, when it is very minutely

squamulose ...” In other words, the ornamentation described by Berkeley

was not the result of age and deterioration, but something observed in young

healthy basidiomes. Redhead did not describe scales from his examination of

the “type” collection, but merely referred to them as “REPORTED squamules,’

suggesting that such scales were reported by Berkeley, but not seen by him.

This strongly suggests that to Berkeley “very minutely squamulose” meant

something other than obviously scaly.

To understand what exactly Berkeley means, it helps to know that he was

a cryptogamist and active microscopist, to whom observing and recording

features best seen with magnification was second nature. As a vicar he likely

also had a good understanding of the English language, with an ability to

use words accurately and precisely. Employing four levels of diminution, he

could not be more emphatic that he is not describing an obvious scaliness like

that seen on scaly-capped sphagnicolous arrhenias (Fic. 3D), but something

much more subtle.

1. He does not say squamous/squamose, but uses the diminutive,

squamulose.

2. He does not stop at squamulose but specifies MINUTELY squamulose.

3. “Minute” is not small, but EXTREMELY small.

4. The diminutive scales of the stipe are not extremely small, but very

extremely small.

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 809

Fic. 7. Lichenomphalia umbellifera: stipe characters. Stipe on Sphagnum, adhering by downy

base: A, B, D-F, M. Pileus hollow: G-K. Pileus impervious or nearly so: G, H. Stipe very minutely

squamulose: L-P. Microscopic view of very minute squamules: Q-S. M courtesy of Michael Wood

and N courtesy of Leif Stridvall.

810... Voitk

Scientific writing is bare and dry. Use of modifiers is not capricious, but

intentionally reserved only to convey important descriptive information.

Thus, in the term “squamulose stipe,’ stipe is taken for granted, and

squamulose becomes the important word, distinguishing this stipe from both

glabrous and squamose ones. It is even less common to modify a modifier,

only justified if the second modifier conveys even more important descriptive

information. In the term “minutely squamulose stipe” the most important

descriptive information is conveyed by “minutely”: the extremely small

size of the stipe adornments serves to distinguish these stipes from merely

squamulose ones.

However, the addition of “very” to exactly the same words that Berkeley

used earlier to describe pileal ornaments, does not serve to distinguish

these stipes from other stipes, but rather to indicate that the ornamentation

is similar on both pileus and stipe, save for size. Size, of course, is relative,

but generally scientific writing relates size of parts to the primary structure

described (unlike romantic poetry, which relates everything to the author).

Hence, very minute stipe squamules are very minute in relation to the size

of the stipe, not the size of Reverend Berkeley. On a stipe described as a little

over 2 mm wide, scales of sphagnicolous arrhenias, which may approach a

width of 0.25-0.5 mm at their base (i.e. 12-25% of the stipe diameter), would

be unlikely to be considered small, let alone extremely small. On the other

hand, stipe adornments of the near-microscopic scale suggested by the above

interpretation of Berkeley’s description would fit. Such adornments have been

described before for the current L. umbellifera. Peck (1893) stated that the

stipe of Omphalia ericetorum (Pers.) S. Lundell [= L. umbellifera] “may be

either glabrous, pruinose or hairy-squamulose’, and Bigelow (1970) described

the stipe of Omphalina ericetorum (Pers.) M. Lange [= L. umbellifera] as

pubescent in his key, and observed that it was “sometimes pubescent above,

if deeply embedded in wet moss”. Perusing Internet images of L. umbellifera,

reveals several with textured stipes. FIGURE 7 shows two examples, one from

California, photo: Michael Wood (Fic. 7M), and one from Sweden, photo:

Leif Stridvall (Fic. 7N).

A search for stipe texture on my photographs proved rewarding. As

suspected at the outset, on photos meant primarily to record collections,

most stipes lacked adequate focus, sufficient resolution, or both, to permit

satisfactory magnification, but magnification and focus enhancement of the

best-focussed stipes confirmed that at close range the texture was irregular,

not smooth (Fic. 7L). To examine the nature of this vestiture, I collected some

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 811

fresh specimens of “normal” (in the sense that their stipes appeared glabrous

to the naked eye) basidiomes. Macro lens photos revealed that their vestment

varied from finely tomentose (Fic. 7P) to wider projecting ornamentation

(Fic. 70). Microscopical examination revealed that the stiptipellis consisted

of longitudinally arranged hyphae. Tufts of 4—6 or more round-ended hyphae,

originating below and parallel to the superficial longitudinal layer, made a

sharp right-angle turn to project through it for a length of 30-50 um or more,

and sometimes then bent to run parallel with the stipe on the outside (Fic.

7Q, longitudinal section, and Fic. 7R,S, transverse section). The observed

adornments fit Berkeley’s words accurately, similar features have been

described by several experienced field mycologists, and—most importantly—

no valid alternate interpretation exists to explain Berkeley’s “very minutely

squamulose” stipe. The only available explanation is that Berkeley was precise

with his use of words and took great pains to say exactly what he meant;

attempts to impose other meaning on these words have proven misleading.

Significantly, Berkeley used different words to describe stipe texture of

sphagnicolous arrhenias. The only other time he used exactly the same words,

“very minutely squamulose” to describe stipe ornamentation was to describe

the stipe of a specimen of Ag. umbelliferus [= L. umbellifera] he found on a

mud wall (Berkeley 1836). Because we know that L. umbellifera does not have

a “scaly” stipe in the common meaning of the term, we now have proof that

Berkeley must mean something other than scaly with those words. Of course,

it does not escape the reader that conspecificity is one possible reason for two

specimens (e.g., Ag. sphagnicola and Ag. umbelliferus in Berkeley’s treatment)

to have similar characters.

This detailed analysis of Berkeley’s words, combined with detailed

morphological examination of the stiptipellis, does not strive to show that

L. umbellifera differs from sphagnicolous arrhenias in this regard. So many

characters in Berkeley’s protologue fit L. umbellifera better than sphagnicolous

arrhenias that there is no need to for lengthy analysis to document additional

differences. The purpose of this detailed analysis is, instead, to provide a

parameter for interpreting Berkeley's “minutely squamulose’ in his description

of the pileus of Ag. sphagnicola.

PILEUS: MINUTELY SQUAMULOSE. Discussion of this character—the very

character from which former misinterpretations of this species stem—was

moved here, because, as mentioned above, knowing what is meant by the

“very minutely squamulose” stipe makes it easy to understand what is meant

by the “minutely squamulose” pileus. Describing surface adornments of both

812... Voitk

stipe and pileus with exactly the same words indicates that the appearance

of the pileus texture is similar to that of the stipe, differing only by a slight

increase in size. Squame is scale, squamule is a little scale, and a minute

squamule is an extremely small little scale. Berkeley describes the stipe

vestiture as a VERY extremely small little scale, i. e. something bordering on

the microscopic, fitting Fic. 7L-Q. Given this standard—and no alternate

demonstration of scale vestiture for stipes of sphagnicolous arrhenias is

available—the extremely small little scale (without “very”) of the pileus fits

the range shown by Fic. 6J—M, which is well out of the range of the scales

on scaly-capped sphagnicolous arrhenias (Fic. 3D-G). Hence, Berkeley's

description of cap ornamentation fits very well with L. umbellifera, e.g.,

Fic. 5M, but presents a significant conflict with its application to a truly scaly

sphagnicolous arrhenias, e.g., Fic. 3C-F.

The appeal of the above review is that all characters in the protologue for

Ag. sphagnicola can be verified in specimens of L. umbellifera—including

the stipe and pileus vestiture, something that has eluded previous attempts.

Redhead’s 1979 and Kuyper’s 1995 opinions are confirmed: the protologue

of Ag. sphagnicola fits better with L. umbellifera than with scaly-capped

sphagnicolous arrhenias. It only remains to assess whether acting on this

would upset the nomenclatural stability of established practice.

Current and traditional application

Redhead & Kuyper (1987) referred to Moser’s Omphalina sphagnicola

(Moser 1967), when deciding to maintain “current and traditional application”

of the name sphagnicola. Moser’s usage may have been current in central

Europe twenty years earlier, but certainly not universally so. By choosing a

new name, Peck (1873) became the first to reject Ag. sphagnicola as a fitting

name for a scaly sphagnicolous Arrhenia. In the margin of his early hand-

written notes of Ag. gerardianus, he wrote, “Is this Ag. affricatus” (Fic. 8),

clearly indicating that he considered earlier descriptions and names—surely

including Ag. sphagnicola—before concluding his to be an undescribed

species. Bigelow espoused the use of Ag. gerardianus for a scaly sphagnicolous

Arrhenia, thereby enrolling more adherents to this practice. Redhead’s 1979

declaration that Ag. sphagnicola was a later synonym for G. ericetorum

[= L. umbellifera] persuaded more mycologists to abandon applying

Ag. sphagnicola to sphagnicolous arrhenias. After Kuyper (1995) examined

the protologue of Ag. sphagnicola, synonymized it with Phytoconis

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 813

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Fic. 8. Photo of Peck’s handwritten note made of the syntype from Sand Lake of Agaricus

gerardianus at NYS. The specimen has been missing, but the note remains. Note margin note—

“Is it Ag. affricatus” —proof that Peck considered older names and descriptions, before deciding this

was a new and hitherto undescribed species.

ericetorum (Pers.) Redhead & Kuyper, and applied O. gerardiana to scaly-

capped sphagnicolous arrhenias, even more mycologists followed this

usage. Significant headway was made after Elborne (2008) accepted Peck’s

Ag. gerardianus as the correct name for scaly-capped sphagnicolous

arrhenias, transferring it to Arrhenia. Favre (1948) had followed Lange

and used O. sphagnicola for scaly-capped arrhenias, synonymizing it with

O. gerardiana. He pointed out that Singer also supported this but stated that

Singer synonymized it with O. philonotis instead. Such inconsistency in the

interpretation of Ag. sphagnicola among authors applying it to scaly-capped

arrhenias may have contributed further to the increasing support to abandon

the application of Ag. sphagnicola to sphagnicolous arrhenias.

Although it is difficult to measure “current and traditional application”

accurately, changes over the last four decades suggest that synonymizing

Ag. sphagnicola with the current L. umbellifera, will be no more disruptive than

applying it to a scaly-capped Arrhenia. In such a situation a decision made in

accordance with the Code has the advantage of more effective contribution to

future stability.

814 ... Voitk

Agaricus sphagnicola Berk., The English Flora, vol. 5(2): 67. 1836.

Type: Holotype: unavailable. Neotype (here designated, MBT10004615): UK, Kew

Gardens, K(M)25223, identified as Agaricus sphagnicola by Berkeley and revised as

Gerronema ericetorum by Redhead (1979: p. 339; A study of the sphagnicolous fleshy

Basidiomycetes in the eastern sections of the Canadian boreal forest. PhD thesis,

University of Toronto).

= Lichenomphalia umbellifera (L.) Redhead & al

ComMENT: To avoid further debate and stabilize nomenclature, the name is

neotypified with a specimen identified as L. umbellifera by Redhead (1979).

This synonymy agrees with Kuyper’s opinion, confirmed here, and is in keeping

with the direction of current practice over the last four decades.

Discussion

The universe with all its contents, including various fungi, is “real” and

can be “observed” (Mann 2012). Such observations are fact. In contrast, our

interpretation of the world around us, even if based on observed fact, is not

fact, but opinion. Taxonomy, our system of ranking organisms based on their

presumed evolutional relationship, is not a fact found in the universe, but a

philosophical construct made by us in an effort to understand and explain

the world we observe. Taxonomy is in constant flux, ever changing as new

information becomes available. Initial ranking was based on macroscopic

appearance. Once observations of distribution, habitat, substrate, and ecology

were seen to separate some organisms from others, ranking changed in some

cases. The ability to observe microscopic morphology brought about significant

taxonomic changes. Ability to analyze chemical composition, and performance

of mating studies contributed to further ranking changes. Our recent ability

to determine DNA sequences cheaply and accurately has introduced a major

revolution in taxonomy. Among other things, molecular studies have allowed

us to recognize species complexes (several separate species, previously

considered one because they look alike), e.g., morels (Richard & al. 2015), on

the one hand, and plastic species (“true” single species, previously suspected

to be several because of their protean nature), e.g., L. umbellifera (Geml & al.

2012) on the other.

Species, the lowest ranked units of taxonomy, are identified by unique

epithets, which stay the same, regardless of how their taxonomic position

may change. One of the main reasons nomenclature has been successful

is that its epithets replace the diverse regionalism of common names with a

stable nomenclature, fixed across the globe by typification governed by the

Code. Lack of typification permits various interpretations of descriptions and

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 815

illustrations by different authors, at times leading to a situation no better that

the earlier common names. A few years back, within one week I got two letters

about one species that began, “Our Nordic interpretation of species X...” and

“The Iberian interpretation of species X....” Needless to say, species X was a

non-typified species. The more complicated our system becomes, and the more

rapidly knowledge that alters taxonomy advances, the more important it is that

names identifying individual organisms remain firmly fixed. In the interests of

more reliable data, many scholarly publications no longer accept ITS analysis

alone, but require multilocus analysis of sequenced species, yet these same

publications regularly present works treating untypified species. With due

respect I submit that it is not the lack of DNA that makes a species concept a

mere rumour, but the lack of typification.

The Code has been established and continuously refined to help new

typification where type material is unavailable. Determining correct application

of names requires interpreting old descriptions, not always easy because these are

often brief, vague and ambiguous (e.g., the epithet icmadophilus). Fortunately,

the descriptions of Ag. cespitosus, Ag. oniscus, and Ag. sphagnicola are lucid, and

the first two have readily identifiable original material suitable for typification.

Words need very careful interpretation, in context, as demonstrated by the case

of Ag. sphagnicola to seek an accurate fit for typification. The use of old names is

desirable: it respects earlier workers and reduces the clutter of orphaned names

clamouring for adoption. Placing names in synonymy also reduces clutter and

seems more respectful than their willful use contrary to the original author's

intent.

The treatment of our three species can be capsulated as follows:

1. Agaricus cespitosus. This is a later synonym for the current L. umbellifera.

No other species has a convincing or congruent fit with the protologue and

lectotype, an opinion verified by an expert panel.

2. Agaricus oniscus. Fries expressed uncommon certainty that the species is

synonymous with Ag. cespitosus, citing its illustration as the only de facto

lectotype. We have formalized his intent by declaring it so. The current

(and retroactive) Code condones change in the description of a species—

whether by the author or another—only if it is supported by the original

material. In this case Fries’s later description and illustration (Fic. 3C) must

be dismissed because one is not supported by the protologue, and the other

is entirely dissimilar from the lectotype (Fic. 2C). Because Ag. cespitosus is a

later name for the current L. umbellifera, Ag. oniscus is also a later synonym

for the current L. umbellifera.

816 ... Voitk

3. Agaricus sphagnicola. The protologue does not support applying this

name to scaly-capped sphagnicolous arrhenias but fits with the current

L. umbellifera. This view concurs with two past assessments (Redhead 1979,

Kuyper 1995), and is strengthened by the fact that at least one specimen

identified by Berkeley as Ag. sphagnicola has also been identified as the

current L. umbellifera (Redhead 1979). Agaricus sphagnicola is another later

synonym for the current L. umbellifera.

To prevent undue disruption, the Code provides for a mechanism to fix

names in accordance with past practice. However, if this has been diverse and

inconsistent, as with Ag. oniscus and Ag. sphagnicola, stability is best served

by fixing names with the best match to the protologue and original material

that avoids conflict. Conserving a name erroneously applied only for a segment

of its application at the expense of displacing names of sequence-confirmed

synonyms with better fit offers little advantage to stability, especially over

solutions that select names fitting with the original material. A poor fit accepted

to-day may become tomorrow’s invitation for retypification, thus serving short

term stability only. No doubt future technologies will continue to refine the

accuracy of our nomenclature and taxonomy, but familiarity with the species is

unlikely to go out of fashion, whatever its nom du jour.

Acknowledgments

I thank Scott Redhead for helpful discussion about these taxa over the years.

Scott helped me to grow from a tyro, unaware that these organisms existed, to

somebody who disagrees with him about them—dquite an achievement! I thank

Irmgard Krisai-Greilhuber and Machiel Noordeloos for help with some reference

material, Konstanze Bensch with precision of type material, and Teuvo Ahti for help

with cryptic meanings in old Botanical Latin. I thank Michael Burzynski, Bryce

Kendrick, Anita & Leif Stridvall, Michael Wood, and Wikipedia for permission to

use their photographs, Asa Kruys for searching UPS for material and the photo

of the Quélet collection and UPS for permission to use it, Karl-Henrik Larsson

for information about the specimen declared epitype for Ag. oniscus, Michaela

Schmull and Gretchen Wade of FH, Laura Biscoe of NY, and Amy Morgan and the

Special Collections, USDA National Agricultural Library, for helping locate Bolton’s

illustrations, as well as the libraries or herbaria mentioned for permission to use

them, Isabella Vallette, Michelle Price, and the Conservatoire et Jardin botaniques

de Genéve for permission to use Jeanne Favre's aquarelles, and Henning Knudsen

for the image of Jacob E. Lange’ illustration and Lene Lange for permission to use

it. I am very grateful to Jozsef Geml, Gro Gulden, Pierre-Arthur Moreau, Esteri

Ohenoja and Anna Ronikier for their willingness to participate in the identification

Agaricus caespitosus, Ag. oniscus, and Ag. sphagnicola typified ... 817

panel. I am deeply indebted to Henning Knudsen, Robert Lticking, Tom May,

Scott Redhead, and Rytas Vilgalys for valuable comments, insights and discussion

related to parts of this material, and rush to add that thanking reviewers by name

does not imply their approval of the content. Special thanks are owed to Pierre-

Arthur Moreau and Greg Thorn for critical formal review of the published MS,

resulting in significant improvement of this discourse, and Shaun Pennycook and

Lorelei Norvell of Mycotaxon for unstinting use of their skill in converting maul-

hewn lumber into cabinetry.

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MY COTAXON

October-December 2021— Volume 136, pp. 819-830

https://doi.org/10.5248/136.819

New records from Turkey: Cortinarius barbatus,

C. osmophorus, & C. saturninus

ISMAIL ACAR

Department of Organic Agriculture, Baskale Vocational High School,

Van Yiiztincii Yil University, 65080 Van, Turkey

CORRESPONDENCE TO: iacar2011@gmail.com

AxsstRAcT—Three Cortinarius species, C. barbatus, C. osmophorus, and C. Saturninus,

have been identified as new records for the Turkish mycota based on macro- and

micromorphological characters, and their identifications were supported by molecular

(rDNA ITS) phylogenetic data. The Turkish specimens are illustrated, and their diagnostic

characters are compared with published descriptions of the same species and related taxa.

Key worps— Agaricales, Cortinariaceae, Eastern Anatolia, Bingol, Bitlis, Hakkari

Introduction

Cortinarius (Pers.) Gray is a taxonomically complex genus of

Cortinariaceae that shows a high degree of morphological variability

(Liimatainen 2013). Morphological characters include a convex (often

umbonate) smooth pileus, emarginate to adnate lamellae, brown to rusty

brown ornamented basidiospores, and a cobweb-like partial veil attaching

stipe and pileus (Ammirati & al. 1985, Hoiland & Holst-Jensen 2000). The

two most diagnostic characters of the genus are the presence of a cortina

(partial veil) between the pileus and the stipe and rusty brown spores (Kirk

& al. 2008, Uzun & al. 2013).

Cortinarius is the largest genus of ectomycorrhizal fungi, distributed

worldwide in temperate areas, coniferous forests, and arctic and alpine

tundra (Brandrud & al. 1990-2018). Many Cortinarius species are endemic,

probably due to climatic and edaphic factors (Garnica & al. 2005; Stefani

820 ... Acar

& al. 2014). The many cryptic taxa within the genus complicate species

identification (Niskanen & al. 2013). The varying interpretations of

morphological characters among mycologists often cause disagreements over

species concepts (Liimatainen 2013), which has produced a proliferation of

synonymous species names within the genus.

Most researchers have used molecular markers to support their species

identification (Garnica & al. 2003, 2005, 2009; Peintner & al. 2003; Froslev

& al. 2005, 2007; Ortega & al. 2008; Liimatainen & Niskanen 2013; Stensrud

& al. 2014; Stefani & al. 2014; Liimatainen & al. 2017; Sesli & Liimatainen

2018). Garnica & al. (2005) and Fr@slev & al. (2007) showed that the rDNA

ITS region is a particularly useful marker for species level identification for

Cortinarius.

The literature (Sesli & Denchev 2014; Akata & al. 2015; Giingér & al. 2015;

Sesli & Moreau 2015; Sesli & al. 2015, 2016; Sesli 2018; Sesli & Liimatainen

2018; Acar & Kalmer 2018; Kalmer & al. 2019) has reported 116 Cortinarius

species from Turkey.

Three additional Cortinarius species—C. barbatus, C. osmophorus,

C. Saturninus—reported as new for Turkey and supported by morphological

characters and molecular data are described below.

Materials & methods

Taxon sampling and morphological studies

Cortinarius samples were collected from Turkey’s Bitlis, Hakkari, and Bingél

provinces in 2018. During fieldwork, specimens were photographed with a Canon

(EOS 60D) camera equipped with a Tokina 100 mm macro lens. Macroscopic

characters (pileus, stipe, lamellae, and cortina) were recorded from fresh materials.

Measurements were made of at least 50 spores and 20 basidia and cheilocystidia

from two samples of each specimen. Sections were mounted in distilled water, congo

red, and 3% KOH solution for examination of spores, basidia, basidiospores, and

marginal cells under a Leica EZ4 stereo microscope and tissues under a Leica DM500

research microscope. Measurements were made using the Leica Application Suite

(version 3.2.0) programme and identified according to Cortinarius literature (Lange

1938; Zerova & al. 1979; Arnolds & Kuyper 1995; Phillips 2006; Froslev & Jeppesen

2008; Brandrud & al. 2012, 2014; Liimatainen & al. 2014; Soop 2018). Specimens

were deposited in the Fungarium of Van Yuiztincii Yil University (VANF).

Molecular studies

Genomic DNA was extracted from dried basidiomata using the CTAB

method (Doyle & Doyle 1987). The purity and quantity of extracted DNA were

determined by using NanoDrop2000c UV-Vis Spectrophotometer and 0.8%

Cortinarius spp. new for Turkey ... 821

Fic. 1. Cortinarius barbatus (VANF Acar1076): a, b. Basidiocarp; c, d. Basidiospores;

e, f. Basidia; g, h. Marginal cells; i, k. Pileipellis. (d, f, h, k in Congo Red).

822 ... Acar

agarose gel electrophoresis. DNA was amplified in 25 ul volume mixtures containing

genomic DNA (10 ng/ul), 10x PCR Buffer, MgCl, (25 mM), dNTP mixture (10

mM), selected primer pair (10 uM), Taq polymerase (5u/ul) and sterile water.

The ITS (ITS1-5.8S-ITS2) region was amplified using primer pairs N-nc18S10

5’AGGAGAAGTCGTAACAAG3’ and C26A 5’GTTTCTTTTCCTCCGCT3’ (Wen & Zimmer

1996). The amplicons were run in a 1% agarose gel and visualized by staining with

Gelred dye. Positive reactions were sequenced with forward and reverse PCR primers

in an ABI 3730XL automated sequencer, and the resulting sequences were assembled

and edited using Alibee Multiple Alignment 3.0 software from the GeneBee website

(www.genebee.msu.su/genebee.html). Ambiguous sites were checked manually and

corrected through strand comparison. The newly generated ITS sequences were

deposited in GenBank.

Phylogenetic results

The amplified ITS region was approximately 650 bp long and encompassed

the complete ITS1, 5.8S and ITS2 subregions. Basic Local Alignment Search

Tool (BLAST) (Altschul & al. 1997) analysis was performed using the National

Center for Biotechnology Information (NCBI) database. Identity values of

our three specimens were: C. barbatus 99.14%; C. saturninus 99.78%; and

C. osmophorus 100%.

Taxonomy

Cortinarius barbatus (Batsch) Melot, Doc. Mycol. 20(77): 94 (1989) Fic. 1

PiLEus 20-50 mm, hemispherical when young, later convex to plane, sticky,

glabrous, white to ivory with a white margin. LAMELLAE cream colored when

young, later greyish cream brown and brown colored when mature. STIPE

35-70 x 5-10 mm, cylindrical to ventricose-fusiform and rooting, the aerial

portion is yellowish brown over a white ground, the subterranean portion is

white, longitudinally fibrillose, weakly viscid.

BASIDIOSPORES 7-9.5(-9.8) x 4-5(-5.7) um, amygdaliform, rather weakly

verrucose, pale straw-colored. Bastp1a 24-35 x 6-8.4 um, cylindrical to

clavate, guttulate, hyaline. MARGINAL CELLS 14.5-23 x 5.2-7 um, clavate,

hyaline. PILEIPELLIS hyphae <27 um diam, clamp connections absent.

SPECIMEN EXAMINED: TURKEY, HakkArt, Semdinli, Ovec village, 37.3718°N

44.4762°E, 1484 m, under Populus and Quercus sp., 15.10.2018, I. Acar (VANF Acar1076;

GenBank MN197665).

COMMENTS—Cortinarius barbatus is characterized by its white cap and stipe,

bitter flesh, cream-greyish gills, and amygdaliform spores. According to Soop

(2018), key characters of C. barbatus are its white cap and bitter flesh, which

our material also possesses.

Cortinarius spp. new for Turkey ... 823

Fic. 2. Cortinarius osmophorus (VANF Acar1133): a, b. Basidiocarps; c, d. Basidiospores;

f. Basidia; g, h. Marginal cells; i, k. Pileipellis. (d, f, h, k in Congo Red).

824 ... Acar

Cortinarius barbatus is morphologically similar to C. alboamarescens Kytov.

& al. and C. emollitoides Bidaud & al., but the three taxa are molecularly distinct.

Cortinarius osmophorus PD. Orton, Trans. Brit. Mycol. Soc. 43(2): 210 (1960) Fie. 2

PiLEus 50-90 mm, hemispherical to convex when young, then expanded,

curved when mature, initially light yellow, whitish to cream, pale ochraceous

yellow when young, ochraceous to brownish when mature, surface viscous,

without veil remnants. OpourR strong, sweetish (like orange blossoms).

LAMELLAE greyish white, then brown colored. Stipe 40-70 x 7-17 mm,

whitish, yellowish brown, longitudinally brown fibrillose due to cortina, the

base abruptly marginate and covered with a whitish veil at the margin.

BASIDIOSPORES 9-11(-13) x (4.6—)5-6(—7) um, amygdaliform to citriform or

ellipsoid, coarsely verrucose. BAstp1A 26-40 x 6.5-9.3 um, hyaline, cylindrical

to clavate. MARGINAL CELLS 18-30 x 5-10 um, mainly clavate, sometimes

cylindrical or lageniform. PILEIPELLIS hyphae <8 um diam, clamp connections

present.

SPECIMEN EXAMINED: TURKEY, BINGOL, Geng, Tarlabasi village, 38.6981°N 40.490°E,

1165 m, under Populus and Quercus sp., 03.12.2018, I. Acar (VANF Acar1113; GenBank

MN197666).

ComMMENTS—Phillips (2006) previously reported Cortinarius osmophorus

from beech woods on chalk, while the collection from Bing6l associated with

Populus and Quercus spp.

Cortinarius osmophorus, which is similar to C. aquilanus T.S. Jeppesen &

Fr@slev, C. cruentipellis Kytov. & al., C. caesiocortinatus Jul. Schaff., and C. talus

Fr., characteristically has a strong orange blossom odor resembling that of

Hebeloma sacchariolens.

Cortinarius saturninus (Fr.) Fr., Epicr. Syst. Mycol.: 306 (1838) Fic. 3

PitEus 30-100 mm, obtusely conic, hemispherical or campanulate

when young, with or without low broad umbo, surface smooth, strongly

hygrophanous, pale ochraceous, brown to reddish brown, chestnut-brown with

a purple to violet tinge, centre sometimes red-brown, silvery shining, margin

covered with white fragments and fibrils. LAMELLAE rather broadly adnate,

cinnamon to purple-brown, then ocher brown, often with a violet tinge, edge

paler. Stree 30-80 x 7-18 mm, cylindrical, often slender, swollen towards

base to subbulbous, silvery whitish to greyish violet, apex violet, covered with

longitudinal white fibrils.

BASIDIOSPORES 7.5-11.7 x 4-6 um, oblong, elliptic, amygdaliform,

moderately to strongly verrucose. BasipDIA 25-37 x 5-10 um, hyaline, cylindrical

Cortinarius spp. new for Turkey ... 825

oe kth ee Poe wa

~") oe

Fic. 3. Cortinarius saturninus (VANF Acar1055): a, b. Basidiocarps; c, d. Basidiospores;

e, f. Basidia; g, h. Marginal cells; i-j. Pileipellis (d, f, h, j in Congo Red).

826 ... Acar

to clavate, 4 spored. MARGINAL CELLS 17-25 x 4—7.5 um, cylindrical to clavate.

PILEIPELLIS hyphae <18 um diam, clamp connections absent.

SPECIMEN EXAMINED: TURKEY, BITLIs, Kuzgunkuran gateway, 38.3644°N 42.7561°E,

1938 m, roadside, under Salix sp. trees, 15.05.2018, I. Acar (VANF Acar1055; GenBank

MN197667).

COMMENTS—Cortinarius saturninus is characterized by persistent veil

remnants on the stipe and strongly verrucose spores. Lange (1938) who reported

C. saturninus in Flora Agaricina, describes material that differs slightly [mostly

ellipsoid spores] from our material due to the different ecological conditions.

Our material is highly compatible with the descriptions by Phillips (2006) and

Soop (2018).

Cortinarius saturninus, which is highly variable in the field, may easily

be confused with C. confirmatus Rob. Henry, C. cyprinus Bidaud & al., and

C. imbutus Fr., which can occur in similar habitats.

Discussion

Cortinarius barbatus belongs to C. sect. Vibratiles (in C. subg. Myxacium)

and characterized by white pileus and stipe, bitter flesh, and amygdaloid spores

(Soop 2018; Soop & al. 2019). Cortinarius sect. Vibratiles is characterized by

medium-small size basidiomes, a viscous pileus and stipe, and bitter context.

Phylogenetically C. barbatus appears closely related to C. alboamarescens and

C. emollitoides. Determining C. barbatus and C. alboamarescens to species

can be difficult because of the morphological similarities. ITS sequence

differences separated C. barbatus (our sample) from C. alboamarescens with

fewer than 10 substitutions and indel positions. Cortinarius alboamarescens is

characterized by very small almost subglobose spores and is the only known

white species of the section that occurs in coniferous forests (Ariyawansa &

al. 2015). Cortinarius emollitoides is distinguished from C. barbatus by its

viscid yellow to yellow-brown pileus and less bitter taste.

Phylogenies by Garnica & al. (2005) and Froslev & al. (2007) place

C. osmophorus in C. sect. Calochroi (of C. subg. Phlegmacium). Species in

this section are characterized by basidiomes with an abruptly marginate

stipe base, well-developed gelatinous pileus, and KOH-reactive pigments

(Garnica & al. 2009). Cortinarius osmophorus is characterized by the sticky,

pale yellowto brownish pileus, abruptly marginate stipe, ornamented spores,

and strong odour. The species grows mainly in Fagus forests in the Southern

hemisphere (Phillips 2006, Froslev & al. 2006, Soop 2018). Macroscopically,

C. osmophorus resembles C. aquilanus. However, C. aquilanus, which grows

Cortinarius spp. new for Turkey ... 827

under Fagus sylvatica, is distinguished from C. osmophorus by its fairly sparse

veil remnants and yellowish spots on the stipe (Froslev & Jeppesen 2008).

Additionally, C. caesiocortinatus and C. cruentipellis are separated from

C. osmophorus by their purple-colored lamellae (Soop 2018).

Cortinarius saturninus, which belongs to C. sect. Saturnini (Liimatainen

& al. 2017), differs from related species by its persistent veil remnants on

the stipe and strongly verrucose spores. The species may be confused with

C. imbutus, C. cyprinus, and C. confirmatus, also representatives of Cortinarius

subg. Telamonia, in the field because they can occur in similar habitats.

Lilac-greyish lamellae and abundant veil remnants on the stipe help identify

C. imbutus. Liimatainen & al. (2017), characterize C. confirmatus by by one

of the following character combinations: (1) absence of veil remnants on the

stipe and not fruiting in dense clusters, (2) abundant veil remnants on the

stipe and densely fruiting under Populus alba, or (3) a reddish tinted pileus

and densely fruiting under Betula pendula. Phylogenetically, C. saturninus

(our sample) and its sister species C. cyprinus differ by 3 substitutions and

2 indels.

Both morphological and molecular analyses support the identification of

the newly recorded Cortinarius species in the current study. The addition

of these three additional species increases the total of reported Cortinarius

species in Turkey to 119.

Acknowledgements

I would like to thank to Research Assistant Aysenur Kalmer for useful comments

regarding the molecular section. I also thank Editor-in Chief Norvell and the

pre-submission reviewers Dr Balint Dima (Edtvés Lorand University, Budapest,

Hungary) and Dr Abdullah Kaya (Karamanoglu Mehmetbey University, Turkey) for

their valuable suggestions.

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MY COTAXON

October-December 2021— Volume 136, pp. 831-840

https://doi.org/10.5248/136.831

Notes on lichenized fungi of chroodiscoid Thelotremataceae

from China

ZE-FENG JIA , MIN L1, Yu-Ru Fu, JING Pu

College of Life Sciences, Liaocheng University, Liaocheng, 252059, China

" CORRESPONDENCE TO: Zffia2008@163.com

AxBstTRAcT—Chinese collections representing eleven species in four genera of lichenized

chroodiscoid Thelotremataceae are reviewed taxonomically. Four species—Astrochapsa

platycarpella, Chapsa patens, C. diploschistoides, C. pulchra—are reported as new records

for China. Each new record is described, illustrated, and discussed in detail. Descriptions

and remarks are also provided for the other seven species previously reported from

China—Astrochapsa mirabilis, Chapsa indica, C. leprocarpa, Chroodiscus argillaceus,

Reimnitzia santensis, Rhabdodiscus asiaticus, and Xalocoa ocellata.

Key worps —Graphidales, Ostropomycetidae, Lecanoromycetes

Introduction

The most recent taxonomic concept of lichen forming Lecanoromycetes

accepts the resurrected family Thelotremataceae Stizenb. in Graphidales

(Ostropomycetidae; Kraichak & al. 2018, Liicking 2019). In Thelotremataceae

ascomata are true apothecia with hemiangiocarpous development, where

crater-like apothecia form their double margin from the cupulate proper

exciple and an outer thallus layer (= thalline margin; Matsumoto 2000,

Frisch 2006). The characteristics of proper exciple, thalline margin, columella

structures, carbonization, and other features determine five types that describe

the considerable differences in apothecial morphology: Chroodiscus-type,

Fibrillitheces-type, Myriotrema-type, Ocellularia-type, and Thelotrema-type

(Frisch 2006). The Chroodiscus-type apothecia, referred to as chroodiscoid by

Hale (1981), are rounded, reniform, angular or star-shaped, and usually large

832 ... Jia & al.

with a distinctly exposed disc surrounded by a jagged, fissured, or lobed (rarely

entire) margin that is erect to recurved or exfoliating (Frisch 2006, Mangold

2008). The chroodiscoid genera in Thelotremataceae are: Acanthotrema,

Astrochapsa Parnmen & al., Asteristion, Chapsa A. Massal., Cruentotrema,

Chroodiscus (Mull. Arg.) Mull. Arg., Enigmotrema, Gyrotrema, Gintarasia,

Myriochapsa, Nitidochapsa, Pseudochapsa, Reimnitzia Kalb, Rhabdodiscus

Vain., and Xalocoa Kraichak & al.

The literature has reported seven species—Astrochapsa mirabilis,

Chapsa indica, C. leprocarpa, Chroodiscus argillaceus, Reimnitzia santensis,

Rhabdodiscus asiaticus, and Xalocoa ocellata—from China (see below).

During our recent research and compilation of FLORA LICHENUM SINICORUM,

additional small genera (e.g., Carbacarthographis, Myriotrema, Schizotrema)

have been described from China. (Jia & al. 2017, Liu & al. 2018, Xu & al. 2016,

Jia & Licking 2020). We report here on specimens collected from Fujian and

Hainan provinces that represent four Astrochapsa and Chapsa species new to

China: A. platycarpella, C. patens, C. diploschistoides, and C. pulchra.

Materials & methods

The specimens are deposited in the Fungarium of College of Life Sciences,

Liaocheng University, China (LCUF). An Olympus SZX16 dissecting microscope

and a Olympus BX53 compound microscope were used for morphological studies.

Measurements were taken from mature cross sections of fruit bodies mounted in

water. Ascospore amyloidity was tested using Lugol’s solution. Spot tests were

performed on the thallus surface with K (10% KOH aqueous solution), C (saturated

aqueous NaOCl solution), and PD (saturated p-phenylenediamine solution in 95%

ethyl alcohol). The lichen substances were detected and identified by thin-layer

chromatography, using solvent C (Culberson & Kristinsson 1970, Culberson 1972,

White & James 1985, Jia & Wei 2016).

Taxonomy (Four new records for China)

Astrochapsa platycarpella (Vain.) Parnmen, Liicking & Lumbsch,

PLoS One 7(12): e51392, 9 (2012) FIG. 1A-C

THALLUS crustose, corticolous, brown-olive, smooth or uneven to warty.

APOTHECIA scattered to aggregate in groups, level with the thallus, angular-

rounded, 0.5-1.5 mm diam. MarGIn thin, lobulate to recurved, sometimes

fused with a white pruinose inner surface. Disc exposed, pale brown to

blackish, with distinct and coarse bluish white pruina. PROPER EXCIPLE

fused, brown at the base; periphysoids <35 um long, free. HyMENIUM clear,

50-100 um high; paraphyses unbranched, straight, 1.5-2 um wide, tips slightly

Chroodiscoid Thelotremataceae in China ... 833

thickened, with brownish granules. Asci clavate, 50-90 x 7-9 um, 8-spored.

Ascosporss hyaline, transversely septate, 4-6-loculate, 15-18 x 4-5 um, thin

walled to slightly thickened at the edges of the lumina, with distal oblong ends

tapering, I-.

CHEMiIstTRY: Thallus K-, C-, PD-; no lichen substances detected by TLC.

SPECIMENS EXAMINED: CHINA. HAINAN PROVINCE, Ledong County, Mt. Jianfengling,

Yulingu. alt. 630 m, on bark, 10-XII-2019, M. Li HN19362, HN19365, HN19368,

HN19377, HN19382 (LCUF).

ECOLOGY & DISTRIBUTION: In China on bark in an evergreen tropical rain

forest. Distributed in Africa, South America, and India (Frisch 2006; Joshi et al.

2018). New to China.

REMARKS: Astrochapsa platycarpella is very close to A. astroidea (Berk. &

Broome) Parnmen & al. but differs in having aggregated apothecia, a bluish

white pruinose disc and white pruinose marginal lobules with solitary to fused

apothecia, grayish pruinose discs, and rarely layered lobules. Mangold (2008)

synonymised Chapsa platycarpella under C. astroidea, but Parnmen & al. (2012)

transferred them both to Astrochapsa as separate species. Chapsa alborosella

(Nyl.) Frisch is also similar but has an ecorticate thallus, dispersed apothecia,

and 5-9-septate ascospores.

Chapsa diploschistoides (Zahlbr.) Frisch, Biblioth. Lichenol. 92: 99 (2006) Fic. 1D-F

THALLUS crustose, corticolous, pale to dark olive-grey or brownish, slightly

uneven to warty. APOTHECIA dispersed, level with the thallus, rounded to

angular, 0.5-1 mm diam. Marcin thick, raised, deeply fissured to lobed,

incurved to upright, often in two or three separate layers. Disc slightly exposed,

pale brown, covered by thin white pruina. PROPER EXCIPLE deeply cupulate,

hyaline to pale brown at the base; periphysoids <20 um long, free. HYMENIUM

clear, 70-90 um high; paraphyses unbranched, straight, 2-2.5 um diam, tips

indistinctly moniliform and often a slight dendroid branching, adspersed with

fine brownish granules. Asci narrowly clavate, 70-85 x 8-12 um, 8-spored.

Ascosporss hyaline, transversely septate, 6-locular, 18-25 x 5.5-6.5 um, thin

walled, with subacute ends, I-.

CuHemistry: Thallus K-, C-, PD-; unknown lichen substances detected by

AHR

SPECIMEN EXAMINED: CHINA. HAINAN PROVINCE, Changjiang County, Mt.

Bawangling, Yajia. alt. 650 m, on bark, 08-VII-2019, Y.H. Ju HN19163 (LCUF).

ECOLOGY & DISTRIBUTION: In southern China on bark in a tropical forest near

a valley waterfall. Also known from South Africa (Frisch 2006). New to China.

834 ... Jia & al.

REMARKS: Chapsa diploschistoides is similar to C. alborosella but differs by its

thicker, fissured to fissured-areolate thallus with distinct warts, a more compact

and glossier thallus surface, and apothecia with an incurved layered margin

having a compact (not white-felty) inner surface. Our Chinese material has

smaller ascospores than that from south Africa noted by Frisch (2006; 20-33

x 6-7 um).

Chapsa patens (Nyl.) Frisch, Biblioth. Lichenol. 92: 111 (2006) FIG. 1G-I

THALLUS crustose, corticolous, greenish grey, often somewhat brownish,

slightly uneven to warty. APOTHECIA dispersed, level with the thallus, rounded

to slightly angular, 1-2.5 mm diam. Maren thick, strongly raised, deeply

fissured to lobed, incurved to upright but usually not strongly recurved in

the apical parts. Disc exposed, pale brown, covered by thick and coarse white

pruina. PROPER EXCIPLE deeply cupulate, hyaline to pale brown at the base;

periphysoids <50 um long, free. HYMENIUM clear, 130-200 um high; paraphyses

unbranched, straight, 2-2.5 um diam, tips strongly moniliform and intensely

adspersed with fine brownish granules. Asci clavate, 100-180 x 25-40 um,

1-spored. Ascosporss hyaline (slightly brownish in age), densely muriform,

80-120 x 20-30 um, thin walled, with rounded ends, with a narrow halo, I-.

CuHEmistTry: Thallus K-, C-, PD-; no lichen substances detected by TLC.

SPECIMENS EXAMINED: CHINA. FUJIAN PROVINCE: Qianzhou City, Mt. Jiuxian,

Huizhaochi. alt. 1540 m, on bark, 25-VII-2019, EY. Liu FJ19127-1, FJ19128-1, FJ19131,

FJ19148-2 (LCUE).

ECOLOGY & DISTRIBUTION: In southern China on bark in a subtropical forest.

Widely distributed in Africa, Japan, Sri Lanka, and India (Matsumoto 2000,

Frisch 2006, Joshi & al. 2012). New to China.

REMARKS: Chapsa patens is similar to C. leprocarpa, which differs by having

a slightly lower hymenium and smaller ascospores (65-85 x 15-23 um; see

Chinese material of C. leprocarpa, below). Hale (1981) and Matsumoto (2000)

described C. patens as having stictic acid, while Frisch (2006) noted that the

lectotype definitely lacks stictic acid. The ascospores of C. patens in our Chinese

collection are slightly smaller than those from Africa (90-158 x 22-35 um;

Frisch 2006) and India (80-125 x 20-35 um; Joshi & al. 2012).

Chapsa pulchra (Mill. Arg.) Mangold,

Flora of Australia 57(Lichens 5): 654 (2009) FIG. 1J-L

THALLUS crustose, corticolous, thin, pale gray to pale grayish-green,

smooth to usually rough. APOTHECIA scattered, level with the thallus,

Chroodiscoid Thelotremataceae in China... 835

4 Py \

( Pit

oS rN \

Fic.1 Astrochapsa platycarpella (LCUF -— Li HN19365): A: habit; B: apothecial section;

C: ascus containing ascospores. Chapsa diploschistoides (LCUF - Ju HN19163): D: habit;

E: apothecial section; F: ascus containing ascospores. Chapsa patens (LCUF - Liu FJ19131):

D: habit; E: apothecial section; F: ascus containing one muriform ascospore. Chapsa pulchra

(LCUF - Li HN19422): G: habit; H: apothecial section; I: ascus containing ascospores.

Scale bars: A, D, G, J = 1 mm; B, E, H, K = 50 um; C, FI, b= 20 um.

836 ... Jia & al.

roundish to irregular, 1-2 mm diam. Marain thick, lobed to eroded, rarely

slightly layered, pruinose, becoming erect to recurved. Disc partly to entirely

visible from surface, pale flesh-colored to grayish, with distinct pruina.

PROPER EXCIPLE fused, brown at the base; periphysoids <30 um long, free.

HyYMENIvM clear, 100-130 um high; paraphyses unbranched, straight, 1.5-2

um diam, tips slightly thickened. Asci clavate, 80-100 x 10-18 um, 6- or

8-spored. Ascospores hyaline, transversely septate, 12—-20-loculate, 30-65

x 5-7 um, thin walled to slightly thickened at the edges of the lumina, with

distal oblong ends tapering, I-.

CuHEmMistTry: Thallus K-, C-, PD-; no lichen substances detected by TLC.

SPECIMEN EXAMINED: CHINA. HAINAN PROVINCE: Ledong County, Mt. Jianfengling,

Yulingu. alt. 630 m, on bark, 10-XII-2019, M. Li HN19422 (LCUF).

ECOLOGY & DISTRIBUTION: In southern China on bark in an evergreen tropical

rain forest. Previously known only from Australia (Mangold 2008). New to

China.

REMARKS: The species is very similar to Chapsa indica, which produces much

larger ascospores with more septa (see below). Another similar species is

Chapsa alborosella (Nyl.) Frisch, which is distinguished by smaller ascospores

with fewer septa (5-9-septate, 17-22 x 3-6 um; Mangold 2008).

Seven other thelotrematoid species known from China

Astrochapsa mirabilis (Zahlbr.) Licking & S. Joshi,

Lichenologist 50(6): 633 (2018)

The species is characterized by a corticate olive-green to greyish green

thallus, angular apothecia with recurved margins, greyish disc covered with

thick white pruina, carbonized exciple, inspersed hymenium, 8-spored asci,

brown muriform 8-10/3-4-loculate ascospores measuring 40-50 x 11-15 um,

and stictic acid.

Reported from Fujian Province of China (Zahlbruckner 1930, Jia & Liicking

2017) and India (Joshi et al. 2018). Pantropical Asia.

Chapsa indica A. Massal., Atti Reale Ist. Venet Sci. Lett. Arti., Sér. 3, 5: 257 (1860).

The species is characterized by the small angular or shortly elongated

apothecia with a usually densely white-felty disc, Chapsa-type paraphyses

and periphysoids, large hyaline transversely septate 18-35-loculate ascospores

(65-100 x 5.5-8 um), and the absence of lichen compounds. Its apothecia,

ascospore type, and lichen compounds are similar to C. pulchra, which differs

by its smaller ascospores (see above).

Chroodiscoid Thelotremataceae in China ... 837

Reported from Hainan Province of China (Xu & al. 2016). Distributed

mainly in Africa (Sierra Leone, Kenya, Mozambique), Asia (India, Sri Lanka,

Borneo, Malaya, Andaman Islands), and Australia (Patwardhan & Nagarkar

1980; Nagarkar & al. 1986, 1988; Frisch 2006; Mangold 2008). Pantropical,

extending to subtropical regions.

Chapsa leprocarpa (Nyl.) Frisch, Biblioth. Lichenol. 92: 108 (2006).

The species is characterized by the chroodiscoid apothecia with lobed and

recurved margins, white heavily pruinose disc, one solitary muriform ascospore

per ascus (60-85 x 15-23 um), and absence of lichen compounds. It is similar

to Chapsa patens, which is distinguished by larger ascospores (see above).

Reported from Guangxi Province of China (Xu & al. 2016). Distributed

mainly in Africa, Sri Lanka, India, Borneo, USA, Brazil, and Australia (Hale

1981, Sipman 1993, Frisch 2006, Mangold 2008). Pantropical, extending to

subtropical regions in both hemispheres.

Chroodiscus argillaceus (Mill. Arg.) Liicking & Papong,

Bryologist 112(1): 154 (2009)

The species is characterized bya grayish green thallus, immersed to erumpent

apothecia with recurved margins, flat opened dark grey discs, 8-spored asci,

colorless ellipsoid 3-septate ascospores (6-10 x 2.5-3 um), and absence of

lichen substances. It is similar to Chroodiscus khaolungensis Papong & Licking,

which has larger ascospores (18-24 um; Papong & al. 2009, Wang & Wei 2018).

Reported from Hainan Province of China (Wang & Wei 2018). Distributed

mainly in Southeast Asia (Papong & al. 2009).

Reimnitzia santensis (Tuck.) Kalb, Mycotaxon 79: 325 (2001)

The species is characterized by a thallus with large columns of calcium oxalate

crystals and abundant isidia, chroodiscoid ascomata, brown submuriform

ascospores (15-25 x 8-12 um), pycnidia with bacilliform conidia, and absence

of lichen substances.

Reported from Zhejiang Province of China (Dou & al. 2019). Pantropical

(Kalb 2001, Frisch 2006, Mangold 2008).

Rhabdodiscus asiaticus (Vain.) Rivas Plata, Licking & Lumbsch,

Taxon 61(6): 1175 (2012)

= Thelotrema formosanum Zahlbr., Repert. Spec. Nov. Regni Veg. 33: 24 (1933)

The species is characterized by its greyish to olive thallus, semi-emergent

apothecia, thick brownish to distinctly carbonized proper exciple, well-

838 ... Jia & al.

developed carbonized columella, 8-spored asci, transversely septate to

indistinctly submuriform hyaline (occasionally pale brownish) ascospores

(10-40 x 7-12 um), and psoromic acid.

Rhabdodiscus asiaticus resembles R. fissus (Mull. Arg.) Vain., which differs

in having smaller brown submuriform ascospores. Aptroot (2004) listed

Thelotrema formosanum as a synonym of Ocellularia asiatica [= R. asiaticus],

noting that its Taiwanese type had been erroneously cited as “Asahina no. 330”

[correctly TNS Asahina F230], and that psoromic acid (TLC; PD+ deep yellow)

was detected from the thallus, the ascospores are I+ violet, and the columella is

conspicuous, simple, and visible from the outside.

Reported from China's Taiwan Province (Zahlbruckner 1933, as Thelotrema

formosanum; Wang-Yang & Lai 1973, as T. formosanum; Aptroot 2004).

Distributed mainly in Vietnam and India (Vainio 1907, Joshi et al. 2018).

Pantropical Asia.

Xalocoa ocellata (Fr.) Kraichak, Liicking & Lumbsch,

Austral. Syst. Bot. 26(6): 472 (2013)

The species is characterized by the greyish to pale yellow thallus, sessile

discoid lecanoroid apothecia (1-2 mm diam.), grayish pruinose disc, thick non-

carbonized proper exciple, 8-spored asci, ellipsoid muriform 3-5/0-2-septate

brown ascospores (15-30 x 7.5-15 um), and norstictic acid.

Xalocoa ocellata resembles Diploschistes cinereocaesius (Sw.) Vain., which

differs by its larger (<3.6 mm diam.) apothecia, and lecanoric acid with traces

of diploschistesic acids.

Reported from Yunnan and Sichuan Provinces (Zahlbruckner 1930, as

Diploschistes ocellatus). This species is cosmopolitan (Kraichak & al. 2013).

Acknowledgments

This study was supported by the National Natural Science Foundation of China

(31750001 & 31270066). The authors are grateful to the presubmission reviewers Prof.

Qiang Ren (State Key Laboratory of Mycology, Institute of Microbiology, CAS) and

Dr. Lulu Zhang (College of Life Sciences, Shandong Normal University, China) for

reading and improving the manuscript.

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MYCOTAXON

October-December 2021—Volume 136, pp. 841-852

https://doi.org/10.5248/136.841

Five new foliicolous micromycete records from Turkey

GOKHAN DoGan’, MAKBULE ERDOGDU’ , ZEKI AYTAG},

ALI IHSAN KARAYEL’, ZEKIYE SULUDERE?, TUGBA ERTUGRUL?

‘Graduate School of Natural and Applied Science,

Kirsehir Ahi Evran University, Bagbas1, Kirsehir, Turkey

*Department of Landscape Architects, Faculty of Agriculture,

Kirsehir Ahi Evran University, Bagbas1, Kirsehir, Turkey

*Department of Biology, Faculty of Science, Gazi University,

Teknikokullar, Ankara, Turkey

* CORRESPONDENCE TO: merdogdu@ahievran.edu.tr

ABSTRACT—Among microfungi collected on foliar spots of vascular plants in Yedigéller

National Park, Bolu Province, Turkey, five species represent new records for Turkey: Ascochyta

daturae, A. euonymi, Mycosphaerella laureolae, Phyllosticta datiscae, and Rhabdospora visci.

Detailed descriptions and photographs of these species are provided.

Key worps—Ascomycota, biodiversity, new host, SEM, taxonomy

Introduction

Microfungi can cause various diseases through leaf infection. Ascochyta

Lib., Mycosphaerella Johanson, Phyllosticta Pers., and Rhabdospora (Durieu

& Mont.) Sacc. are among the genera that include species causing foliar

spots. Some of these foliar-pathogenic fungi show host specificity, while

others can be found on several hosts: e.g., Rhabdospora visci infects only

Viscum album, while Ascochyta daturae is observed on more than one

species in Solanaceae.

Foliar pathogens causing needle or leaf-bound diseases can cause growth

reduction asa result of a decrease in photosynthetic capacity at high levels of

infection (Van der Pas 1981, Manter & al. 2003, Hanso & Drenkhan 2012).

842 ... Dogan & al.

Tree mortality can be seen only in serious cases. By weakening the tree, leaf

diseases can contribute to higher sensitivity to biotic and abiotic stresses

(Bednaiova & al. 2013, Kowalski 2013). Based upon trophic interactions

with the host, different pathogen species can be distinguished; for instance,

while necrotrophic fungi live on dead cells, biotrophic fungi directly derive

carbon and nutrients from living cells (Deacon 1997).

The Yedigdller National Park, located in the northern part of Bolu

Province in the Western Black Sea Region in Turkey, includes seven lakes

and many streams. The park is situated in square A3 of the Davis (1965-

85) grid square system. The climate of the province is oceanic, and it rains

during all seasons.

The research area comprises primarily mixed forest vegetation. At

lower elevations the forest is dominated by Fagus orientalis, Carpinus

orientalis, Quercus spp., Sambucus nigra, Sorbus aucuparia, S. torminalis,

Corylus colurna, Acer campestre, A. platanoides, Cornus mas, Populus

tremula, and Alnus glutinosa. Regions at upper elevations are covered with

Gymnospermae such as Abies nordmanniana subsp. equi-trojani, Pinus

nigra, P. sylvestris, and Taxus baccata. The understory contains shrubs such

as Daphne pontica, Rhododendron ponticum, and Juniperus oxycedrus. The

research area is also rich in marshy habitats, and there are aquatic plants in

the lake (e.g., Potamageton sp., Lemna sp.) and marsh plants surround the

edge of the lake (Typha sp., Carex spp., Lythrum sp.).

Materials & methods

Plant specimens infected with microfungi were collected in the Yedigdller

National Park, Bolu Province, Turkey. Host specimens were prepared following

conventional herbarium techniques. Host plants were identified using the FLorA

OF TURKEY AND EAST AEGEAN ISLANDS (Davis 1965-85). Thin sections prepared

from infected host tissue were examined under a Leica DM E light microscope and

measured from mounts in tap water. Twenty microscopic structures were measured

for each sample. Infected host surfaces were photographed using a Leica EZ4D stereo

microscope. Species were identified using relevant literature (Ascochyta: Vanev

& al. 1997, Mel’nik 2000, Pote¢ & Ruszkiewicz-Michalska 2011; Mycosphaerella:

Ciferri 1956; Phyllosticta: Sydow 1899, Cejp 1965; Rhabdospora: Winter 1883 [as

Septoria], Saccardo 1884 [as Septoria], Diedicke 1914). All examined specimens

were deposited in the Mycological Collection of the Department of Landscape

Architects, Faculty of Agriculture, Kirsehir Ahi Evran University, Kirsehir, Turkey

(AEUT).

Microfungal foliar pathogens new to Turkey ... 843

For scanning electron microscopy (SEM), infected leaves were mounted on

stubs with double-sided tapes. They were coated with gold in Polaron SC 502

Sputter Coater and examined with Jeol JSM 6060 SEM at 5-10 kV in the Faculty of

Science, Gazi University, Ankara, Turkey.

Taxonomy

During our study on microfungi on vascular plants in Yedigéller National

Park in Turkey we collected several microfungal species that cause foliar

, ="

= 4

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Fic. 1. Ascochyta daturae (AEUT GD1376). A. leaf spot (detail) showing pycnidia; B. pycnidia on

leaf (SEM); C. pycnidium, vertical section; D, E. conidia; F. conidium (SEM).

844 ... Dogan & al.

spots. A check of the current literature revealed that among the microfungal

species causing foliar spots determined in this study, Ascochyta daturae,

Ascochyta euonymi, Mycosphaerella laureolae, Phyllosticta datiscae, and

Rhabdospora visci represent new records for Turkey. The list of microfungi

with their descriptions, host plants, habitats, localities, collection dates, and

voucher numbers follow the classification provided by Index Fungorum

(2020) with slight modifications.

Ascochyta daturae Sacc., Michelia 1(2): 163 (1878) FIG. 1

Spots visible on both sides of leaves, generally circular or angular,

5-15 mm diam., sometimes elongated 23-45 x 12-15 mm, pale brown,

margin of spots dark brown. Conrpiomata pycnidial, epiphyllous,

aggregated or scattered, unilocular, globose, globose-depressed, semi-

immersed, 100-190 x 85-140 um, light brown; ostiole circular or sometimes

papillate, 25-30 um diam. Conrp1A cylindrical, oblong, ellipsoid, 1-septate,

rarely aseptate, straight or slightly curved, not or slightly constricted, both

ends rounded, sometimes one end slightly narrowed, 6.4-11 x 2.8-4 um,

guttulate, hyaline.

SPECIMEN EXAMINED— TURKEY, Bo Lu PROVINCE: Yedigéller National Park, near the

Bityik Lake, 40°56’28”N 31°44’51”E, 770 m asl, on living leaves of Atropa belladonna L.

(Solanaceae), 28.06.2018, G. Dogan (AEUT GD1376).

DISTRIBUTION—Bulgaria (Vanev & al. 1997); Canada (Ginns 1986); China

(Tai 1979); Poland (Mulenko & al. 2008); Venezuela (Urtiaga 1986).

Notes: Ascochyta consists of facultative parasites that cause diseases

of many cultivated and wild plants. A few species are known only from

dead parts of their host plants (Melnik 2000). In Turkey, Ascochyta

species are poorly known and not yet intensively studied. Some species

of Ascochyta were reported by Bremer & al. (1947, 1948), Karel (1958),

Gobelez (1964), Parlak & Gucin (1993), Eken (2003), and Kabaktepe &

al. (2019). Bahcecioglu & Kabaktepe (2013) listed 20 Ascochyta species

from Turkey, on 25 plant species (in 21 genera and 16 families). Mel’nik

(2000) synonymised A. atropae Bres. (the only species recorded on Atropa

belladonna) with A. daturae, which has been recorded on living and dry

leaves and other parts of members of Solanaceae. Microscopic features of

our specimen corresponded to those given by Melnik (2000) and Pole¢ &

Ruszkiewicz-Michalska (2011). We observed unicellular conidia on our

specimens similar to those reported by Pote¢ & Ruszkiewicz-Michalska

(2011).

Microfungal foliar pathogens new to Turkey ... 845

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Fig. 2. Ascochyta euonymi (AEUT GD1980). A. leaf spots; B, C. pycnidia on leaf (SEM);

D. pycnidium, vertical section; E. conidia; F. conidia (SEM).

Ascochyta euonymi Pass., Diagn. Funghi Nuovi 5: 11 (1891) FIG. 2

Spots visible on both sides of leaves, mostly circular, 10-15 mm diam.,

whitish, surrounded by a brown border. Conrpromata pycnidial, epiphyllous,

scattered, unilocular, globose, semi-immersed, 60-115 um diam., brown;

ostiole papillate, 27-40 um diam. Conrp1a cylindrical, ellipsoid, lanceolate,

straight, aseptate or 1-septate, not constricted, both ends rounded, sometimes

one end slightly narrowed, 3.8-6.8 x 2-3 um, guttulate, hyaline.

846 ... Dogan & al.

SPECIMEN EXAMINED—TURKEY, BOLU: Yedigéller National Park, 40°53’45’N,

31°40'06”E, 1627 m asl, on living leaves of Euonymus verrucosus Scop. (Celastraceae),

02.08.2019, G. Dogan (AEUT GD 1980).

DISTRIBUTION—Czechoslovakia (Farr & Rossman 2020); Georgia

(Nakhutsrishvili 1986); Italy (Mel'nik 2000).

Notes: Ascochyta euonymi, A. euonymella (Sacc.) Allesch., A. euonymicola

Allesch., and A. oudemansii Sacc. & P. Syd. are reported to infect Euonymus

spp. (Mulenko & al. 2008, Mel’nik 2000). Our specimen is morphologically

similar to Ascochyta euonymias described by previous study (Mel'nik 2000),

but differs in having slightly broader, shorter, 1-2-celled conidia.

Mycosphaerella laureolae (Desm.) Lindau,

Nat. Pflanzenfam. 1(1): 424 (1897) Fic. 3

Spots single or confluent, amphigenous, usually circular, irregular

when confluent, 5-11 mm diam., at first blackish, becoming brown in the

centre, surrrounded by a dark ring. PERITHECIA mostly epiphyllous, rarely

hypophyllous, usually grouped in the centre of the spots, globose, 65-100

um diam., blackish; ostiole papillate, 18-20 um diam. Ascz1 bitunicate,

subcylindric to clavate, attenuate at the base, contracted almost stipitate,

33-50 x 8-10.7 um. AscosporeEs biseriate, oblong, ovate or ellipsoid,

1-septate, straight, not constricted, both ends rounded, 12.4—15.8 x 3.6-4.7

um, guttulate, hyaline.

SPECIMEN EXAMINED— TURKEY, Bo Lu PROVINCE: Yedigéller National Park, Mengen

road separation, 40°56’38”N 31°44’51”E, 741 m asl, on living leaves of Daphne pontica

L. (Thymelaeaceae), 03.05.2019, G. Dogan (AEUT GD1792).

DIsTRIBUTION—Dominican Republic (Ciferri 1956); France, Germany, and

Switzerland (Stevenson 1926); Morocco (Rieuf 1970, as Sphaerella laureolae).

Notes: Mycosphaerella s.lat. is one of the largest groups of ascomycetes

and includes more than 3000 taxa, with species recognized as pathogens or

endophytes of many plants, hyperparasites of other fungi, or saprobes (Crous

2009). Our identification of Mycosphaerella laureolae agrees with other

descriptions of the species with respect to the morphology of perithecia, asci,

and ascospores, the only observable difference being the wider dimensions

of ascospores and the smaller dimensions of asci. Ciferri (1956) described

ascospores measuring 10-15 x 2-3 um and asci measuring 40-50 x 8-10 um;

and Saccardo (1882) described ascospores measuring 12-14 x 3 um and asci

measuring 44 x 8 um (as Sphaerella laureolae). Here we report Daphne pontica

as a new host for Mycosphaerella laureolae.

Microfungal foliar pathogens new to Turkey ... 847

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Fic. 3. Mycosphaerella laureolae (AEUT GD1792). A. leaf spot (detail) showing perithecia;

B. perithecia on leaf (SEM); C. perithecium on leaf (SEM); D. perithecium, vertical section;

E. ascus and ascospores; F, G. ascospores mounted in lactophenol cotton blue.

Phyllosticta datiscae P. Syd., Beibl. Hedwigia 38: (135) (1899) Fic. 4

Spots amphigenous, orbicular to oblong, often with concentric rings,

reaching a diam. of about 1 cm, brown, margin of spots dark brown.

ConripiIoMarTa pycnidial, epiphyllous, scattered, unilocular, globose-depressed,

semi-immersed, 110-135 x 70-100 um, brown; ostiole papillate. Conrp1a

one-celled, oblong, ellipsoid, rounded at both ends, 5.5-6.9 x 2.2-2.7 um,

biguttulate, hyaline.

848 ... Dogan & al.

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Fig. 4. Phyllosticta datiscae (AEUT GD2041). A. leaf spot (detail) showing pycnidia;

B. pycnidium, vertical section; C. conidia; D. conidia (SEM).

SPECIMENS EXAMINED—TURKEY, BoLu PROVINCE: Yedigéller National Park,

40°56’52”N 31°45’02’E, 716 m asl, on living leaves of Datisca cannabina L. (Datiscaceae),

27.06.2018, G. Dogan (AEUT GD1328); Mengen road separation, 40°56’30”N

31°44’40’E, 811 m asl, on living leaves of Datisca cannabina, 03.08.2019, G. Dogan

(AEUT GD2041).

DISTRIBUTION—Germany (Sydow 1899), Czechoslovakia (Cejp 1965).

Notes: Phyllosticta is an important genus of plant pathogenic fungi, causing

leaf spots and various fruit diseases worldwide on a large range of hosts (Aa

& Vanev 2002). Species recognition in Phyllosticta has historically been based

on morphology, culture characters, and host association. Although there have

been several taxonomic revisions and enumerations of species, there is still

considerable confusion when identifying taxa (Wikee & al. 2011). Phyllosticta

datiscae is the only species of Phyllosticta described on Datisca cannabina. The

Turkish specimens agree with the description by Sydow (1899) in leaf spot,

conidiomatal, and conidial morphology, except that the conidiomata are larger

and the conidia smaller; in comparison with the description and illustration by

Cejp (1965), the Turkish conidia were thicker and 1-2-celled.

Microfungal foliar pathogens new to Turkey ... 849

Rhabdospora visci (Bres.) Died.,

Krypt.fl. Brandenburg 9(3): 537 (1914 [“1915”] Fic. 5

LEAF SPOTS amphigenous, raised, circular, 2-4 mm diam. or confluent and

larger, pale brown, surrounded by a reddish-brown border. CONIDIOMATA

pycnidial, mostly hypophyllous, scattered or in groups, unilocular,

subglobose, immersed, later becoming erumpent, 175-350 um diam., dark

brown. Conrp1a filiform, straight or slightly curved, 1-3-septate, non-

20 nm

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Fic. 5. Rhabdospora visci (AEUT GD 2014). A. leaf spot (detail) showing perithecia;

B. perithecium, vertical section; C. conidia; D. ascospores mounted in lactophenol cotton blue;

E, F. conidia (SEM).

850 ... Dogan & al.

constricted at the septum, attenuated both ends, (18.5-)24.6-36 x 2.2-2.9

um, guttulate, hyaline.

SPECIMEN EXAMINED—TURKEY, Botu Province: Yedigéller National Park,

40°93’58”"N 31°75’39’E, 1083 m asl, on living leaves of Viscum album subsp. abietis

(Wiesb.) Abrom. (Santalaceae), 02.08.2019, G. Dogan (AEUT GD 2014).

DisTRIBUTION—Austria/Italy (Winter 1883, as Septoria visci); Germany

(Diedicke 1914); Hungary (Geza & al. 2009, as Septoria visci); Italy (Farr

& Rossman 2020, as Septoria visci); Serbia and Bosnia and Herzegovina

(Stanivukovi¢ & al. 2010, as Septoria visci).

Notes: Biological control of parasites by using plant pathogens has gained

acceptance as a practical, safe and environmentally beneficial management

method applicable to agro-ecosystems (Charudattan 2001). Control of

European mistletoe is a major problem for the forest service in Turkey (Yuksel

& al. 2005). More than 20 microscopic fungi live on European mistletoe, but

only a few of them cause major damage on the plant (Karadzic & al. 2004).

Of these, Rhabdospora visci, which causes leaf spot disease of European

mistletoe, appears to have potential as a biological control agent against of this

semiparasite.

This species was initially described as Septoria visci on leaves of Viscum

album parasitising gymosperm trees in South Tyrol (Winter 1883). It was

later transferred to the genus Rhabdospora by Diedicke (1914). Our specimen

of Rhabdospora visci was morphologically similar to specimens described by

previous studies (Winter 1883, Diedicke 1914) but differed in having slightly

thicker and shorter conidia.

Acknowledgments

The authors thank Meike Piepenbring (Goethe University, Frankfurt am

Main) and Sabeena Aliyarukunju (Jawaharlal Nehru Tropical Botanic Garden and

Research Institute, India) for pre-submission review. This work was supported by

the Scientific and Technological Research Council of Turkey (TUBITAK) (Project

No: 217Z038).

Literature cited

Aa HA van der, Vanev S. 2002. A revision of the species described in Phyllosticta. CBS, Utrecht.

510 p.

Bahcecioglu Z, Kabaktepe $. 2013. Checklist of Ascochyta (Pleosporales Ascomycota) species found

in Turkey. Journal of the Faculty of Forestry, Istanbul University 63(1): 47-49.

Bednarova M, Dvorak M, Janousek J, Jankovsky L. 2013. Other foliar diseases of coniferous trees.

458-487, in: P Gonthier, G Nicolotti (eds). Infectious Forest Diseases. Wallingford, CABI.

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MY COTAXON

October-December 2021— Volume 136, pp. 853-863

https://doi.org/10.5248/136.853

Cribraria lepida, Physarum dictyosporum, P. diderma,

and P. spectabile newly recorded from Turkey

GONUL EROGLU’®

Selcuk University, Science Faculty, Biology Department,

Konya, Turkey

“ CORRESPONDENCE TO: gnleroglu@gmail.com

ABSTRACT—The moist chamber technique was used to obtain Cribraria lepida, Physarum

dictyosporum, and P. diderma from wood or bark at different outdoor localities in Karaman

Province and P. spectabile in Konya Province. These taxa are reported for the first time from

Turkey.

Key worps—Cribrariaceae, Mycetozoa, myxobiota, Physaraceae, slime moulds

Introduction

The myxomycetes, or slime moulds (Mycetozoa or Myxogastria), are a

group of ameboid protists, traditionally studied by mycologists. Martin &

Alexopoulos (1969) considered the myxomycetes to represent fungi, but

at various times they have been classified as animals, plants, or fungi (Ing

1994). Recently, Cavalier-Smith (2013), following zoological nomenclature,

proposed a new classification of the myxomycetes based on evolutionary and

phylogenetic evidence; myxomycetes are currently classified as Myxogastria

in the super-class Amoebozoa and first rank Eumycetozoa. Fruiting bodies

produced by amoeboid organisms are referred to as either sporocarps,

developing from a single amoeboid cell (myxomycetes and protostelids), or

sorocarps, developing from an aggregate of amoeboid cells (dictyostelids)

(Adl & al. 2005). Myxomycetes form a monophyletic group within the

Amoebozoa (Adl & al. 2018, Leontyev & al. 2019). Despite formal classification

as Amoebozoa, myxomycetes will continue to be studied by mycologists, while

854 ... Eroglu

species classification will likely continue to follow the International Code

of Botanical Nomenclature due to the practicality of retaining the system

and difficulty in transitioning to another nomenclatural system (Keller &

Everhart 2010). In this study, www.eumycetozoa.com was used to determine

the nomenclatural status of the genera and www.mycobank.org was used for

systematic information.

Myxomycetes are free-living predators of other eukaryotic protists and

bacteria and have been recorded in every terrestrial habitat investigated

to date (Stephenson & Rojas 2017). In addition to their abundance in the

soil, they are abundant and more commonly studied in the overlying litter

layer, decaying wood, the bark of living trees, or numerous other substrates

associated with various decaying plant materials (Stephenson 2011, Walker

& al 2019). Specimens for study can be obtained as fruiting bodies that have

developed in the field under natural conditions or cultured in the laboratory

(Stephenson & Rojas 2017).

Myxomycete research in Turkey is relatively recent. The first study of

myxomycetes in Turkey was made by Lohwag (1957). Recently, interest in

these extraordinary organisms has increased in Turkey and a total of 286

myxomycete species have been listed from Turkey (Baba & Sevindik 2019).

This report covers Turkish records of four additional myxomycete species.

Materials & methods

During 2013-2018 field surveys were organized for two different projects in

Karaman. The moisture chamber technique, applied to collected samples in the

cultivated mushroom cultivation room, revealed three new records. A fourth

new record was recovered from Pleurotus ostreatus culture logs (Hadim-Konya)

on which different procedures were performed at the Fungarium of the S.U.

Mushroom Application and Research Center. The physical conditions in the

mushroom cultivation rooms are close to the moist chamber culture technique;

temperature, humidity, and ventilation are automatically controlled as described

by Eroglu & al. (2019): 85-90% humidity; 17-22 °C temperature; 12 hours day

and 12 hours night; 10 minutes per hour ventilation; and water vapor fogging for

5 seconds every half hour was automatic in the production room for Pleurotus

cultivation on poplar logs. Myxomycete development requires higher temperatures

(24-25 °C), but the other conditions are suitable.

Materials were examined under the stereo microscope daily and the developing

samples were glued to cardboard with their substrates and left to dry in the

laboratory. The specimens obtained were dried in the mycology laboratory and

identified by myxomycete identification literature (e.g., Martin & Alexopoulos

1969, Neubert & al. 1993). Fungarium numbers were assigned after identification

Cribraria & Physarum spp. new to Turkey... 855

and illustration of each specimen. During the identification procedure, samples

were photographed with an imaging stereo microscope (SM) and alight microscope

(LM). The specimens are deposited in the Mushroom Application and Research

Center Fungarium, Selcuk University, Konya, Turkey (KONF).

Taxonomy

Fic. 1. Cribraria lepida (KONF GE141).

A. Sporocarps (SM); B. Peridial meshes and spores (LM); C. Spores (LM).

Cribraria lepida Meyl., Bull. Soc. Vaud. Sci. Nat. 56: 326. 1927. FIG. 1

Sporocarps gregarious, stalked. Sporotheca nodding, bright violet,

0.25-0.5 mm diam. Stalk dark violet, 2-5 mm long, 6-10 times the diameter

of the sporocarp. Calyculus occupying one-third of the zoothecia, net

bearing, prominent, thickened, pulvinate nodes. Dictydine granules deep

purple, 0.5-1 um. Spores minutely warted, violet in mass, pale lilaceous by

light microscope, (5—)7-8 um diam.

SPECIMENS EXAMINED: TURKEY. KARAMAN PROVINCE: Sariveliler, Civler village,

36.6680°N 32.5830°E, 1380 m, on debris wood of Pinus sp., 14.11.2015 (KONF GE141);

Dumlugéze, 36.5583°N 32.6017°E, 880 m, on debris wood of Pinus sp., 01.07.2016

(KONF GE291); Basyayla, 36.7611°N 32.6694°E, 1594 m, 15.10.2016 (KONF GE403).

CoMMENTS—Cribraria lepida, which can have a color similar to that of

C. violacea Rex, can be distinguished by the thickened nodes of the peridial

reticulum. C. purpurea Schrad. has longer sporangia with a relatively short

stalk and the dark violet granules are 2-3 (-5) um diam. (Nannenga-

Bremekamp 1991). C. violacea has angular, very large nodes. The nodes differ

considerably: flattened in C. violacea and pulvinate in C. lepida (Ramirez-

Ortega & al. 2017). Cribraria lepida has very long sporangial stalk and is

856 ... Eroglu

violet colored while C. purpurea is reddish violet or violaceous pink and has

a shorter stalk.

Cribraria lepida has been reported from Europe, North and South

America, and Asia [Vietnam, Philippines, Western Siberia] (Bernardo & al.

2018, Estrada-Torres & al. 2009, Lado & al. 2011, Martin & Alexopoulos 1969,

Meylan 1927, Neubert & al. 1993, Novozhilov & al. 2017, Redefia-Sontos &

al. 2018, Vlasenko 2015); here it is reported for the first time from Turkey.

Fic. 2. Physarum dictyosporum (KONF GE102).

A. Sporocarps (SM); B. Capillitium and spores (LM); C. Spores (LM).

Physarum dictyosporum G.W. Martin, Brittonia 14: 183. 1962. Fi. 2

Fructification plasmodiocarpous, scattered with rounded and sessile

sporangia, gregarious, 0.4-0.6 mm diam., 0.5-0.6 mm tall, the plamodiocarps

<2.2 mm long, sinuous, very rarely branched, on inconspicuous dark brown

hypothallus. Peridium double, the outer layer a limy crust, white, the inner

peridium membranous, translucent, irregularly dehiscent. Capillitium

abundant, the nodes white, angular and irregular in shape, connected by

inconspicuous threads. Spores black in mass, violaceous brown in transmitted

light, globose, 10-15 um (mostly 10-11 um) diam., the surface markings

banded reticulate, with tall coarse dark warts connected by stout bands.

Plasmodium white.

SPECIMENS EXAMINED: TURKEY. KARAMAN PROVINCE: Bucakkisla, 36.9233°N

33.0278°E, 956 m, on bark of living Pinus sp., 06.06.2014 (KONF GE102); Taskale,

37.1344°N 33.6072°E, 1378 m, on bark of living Juglans sp., 09.11.2014 (KONF GE305).

COMMENTS—Physarum echinosporum Lister also produces white

plasmodiocarpous sporocarps which, however, are laterally compressed, and

Cribraria & Physarum spp. new to Turkey ... 857

not terete as those of P. dictyosporum (Liu & al. 2013). Physarum dictyosporum

has very distinctive spores but superficially resembles the cosmopolitan

Didymium difforme (Pers.) Gray (Ing & Hnatiuk 1981).

Physarum dictyosporum has been reported from North and Central America,

Africa [Seychelles], and Asia [India, Taiwan] (Alexopoulus & Saenz 1975, Farr

1976, Ing & Hnatiuk 1981, Liu & al. 2013, Ranade & al. 2012, Rojas & al. 2018);

here it is reported for the first time from Turkey.

Fic. 3. Physarum diderma (KONF GE86).

A. Sporocarps (SM); B. Capillitium and spores (LM); C. Spores (LM).

Physarum diderma Rostaf., Sluzowce Monogr.: 110. 1875 [“1874”]. FIG. 3

Sporangia clustered, sessile or narrowly adnate, globose, pulvinate,

about 1 mm. diam., snow-white. Peridium double, the outer layer dense,

fragile, thick, calcareous, the inner delicate, translucent, membranous.

Capillitium abundant, the nodes white, angular, sometimes uniting to form a

pseudocolumella. Spores black in mass, purplish-brown in transmitted light,

rough, 10-12 um diam.

SPECIMENS EXAMINED: TURKEY. KARAMAN PROVINCE: Kazimkarabekir, 37.205°N

32.9469°E, 1154 m, on bark of living Quercus sp., 17.05.2013 (KONF GE12); Bucakkisla,

near Goksu river, 36.9508°N 33.0203°E, 421 m, on debris wood of Pinus sp., 06.06.2014

(KONF GE86).

CoMMENTS—Physarum diderma can be confused with P. didermoides (Pers.)

Rostaf., P. spectabile, and P. bitectum G. Lister. The sporocarps of P. didermoides

are sessile or stalked; when membranous stalked, the stalk often appears like

a flattened extension of the hypothallus. Physarum bitectum sporocarps are

slightly compressed laterally contrasting with the sessile globose sporocarps

858 ... Eroglu

of P. diderma. The capillitia of P. didermoides and P. licheniforme (Szabo

ex Schwein.) Lado are small and rounded, in contrast to the large angular

capillitium in P. diderma (Poulain & al. 2011).

Physarum diderma has been reported from Europe, North America, and

Asia [Israel, India] (Eliasson & Adamonyte 2009, Harling 1952, Martin 1949,

Novozhilov & al. 2006, Ukkola & al. 1996, Zoll & Stephenson 2015); here it is

reported for the first time from Turkey.

Fic. 4. Physarum spectabile (KONF GE747).

A. Sporocarps (SM); B. Capillitium and spores (LM); C: Spores (LM).

Physarum spectabile Nann.-Bremen., Lado & G. Moreno,

Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 87(1): 91. 1984. Fic. 4

Sporangia sessile; crowded or scattered, sometimes heaped; flattened,

pulvinate, on wide base; sporangia 0.5-0.7 mm diam., plasmodiocarps

1-2.5 x 0.5-0.7 mm, white. Peridium single, membranous, white or ash-

gray, encrusted with granular white lime in upper part. Hypothallus

membranous, translucent, inconspicuous. Stalk and columella absent.

Capillitium comprising a small-meshed net of colorless fine tubes with many

rounded, (sometimes oblong and branched) lime nodes, sometimes forming

pseudocolumella. Spores dark brown or almost black in mass; dark purplish

brown in transmitted light, angular in optical section, with pale narrow lines,

12-14 um diam., densely ornamented regularly distributed warts.

SPECIMEN EXAMINED: TURKEY. KonyA PROVINCE: Hadim, Egiste stream, 35.1083°N

32.4558°E, 1081 m, on bark of Populus sp. culture log of Pleurotus ostreatus, 18.10.2017

(KONF GE747); Egiste stream, 37.10944°N 32.4575°E, 1095 m, on bark of Populus sp.

culture log of P. ostreatus, 18.10.2017 (KONF GE755).

Cribraria & Physarum spp. new to Turkey ... 859

COMMENTS—Physarum spectabile has very variable characters and may be

either sessile or have a well-defined calcareous stalk. The species can be

confused with P. straminipes Lister, which has a non—-calcareous membranous

extension of the hypothallus and not a stalk. Even when P. spectabile is sessile,

the narrow pale bands on the usually angular spore separate P. straminipes from

P. atacamense D. Wrigley & al. and Badhamia melanospora Speg. (Nannenga-

Bremekamp & al. 1984, Lado & al. 2007, Estrada-Torres & al. 2009, Wrigley

de Basanta & al. 2012, Gmoshinskiy & al. 2017). Badhamia melanospora has

a completely different badhamioid capillitium with calcareous tubes in a

network, not limeless tubes connecting small round lime nodes (Castillo & al.

1996, Moreno & Oltra 2010, Gmoshinskiy & al. 2017).

Physarum spectabile has been reported from Europe, North and South

America, and Africa (Estrada-Torres & al. 2009, Gmoshinskiy & al. 2017, Kylin

& al. 2013, Lado & al. 2007, Lizarraga & al. 2015, Nannenga-Bremekamp & al.

1984, Ndiritu & al. 2009, Oltra & Tejedor 2005, Xavier de Lima & Cavalcanti

2017, Yatsiuk & al. 2017); here it is reported for the first time from Turkey and

from Asia.

Discussion

Cribraria was proposed by Persoon (1794); and Cribraria species are

characterized by dictydine granules, peridium remaining as a net with

without nodes, and sporangiate fruiting bodies consisting of stalked

sporangia with or without basal cups (Liu & Chang 2007). According to

Martin (1949), Martin & Alexopoulos (1969), and Nannenga-Bremekamp

(1991), plasmodic granules, the lack of a capillitium, and the persistent

surface net make Cribraria species easy to recognize (Ramirez—Ortega & al.

2009). The plasmodic granules of Cribraria are also referred to as dictydine

granules (Martin & Alexopoulos 1969, Keller & Braun 1999, Stephenson

2003), calcareous dictydine granules (Ing 1999), lime globules (Nannenga-

Bremekamp 1991) or calcic granules (Discover Life 2020).

Physarum was also proposed by Persoon (1794). The name was coined

based on the bubble-like appearance of the sporangium (Ranade & Ranadive

2016). The single most important character in Physarum is the presence

of lime (calcium carbonate) deposits that may occur in the peridium,

capillitium, or sporothecal stalk (Stephenson & Stempen 2000). With about

149 taxa worldwide, Physarum is the largest myxomycete genus (Lado 2020)

and the most widely known genus among the myxomycetes (Hoppe & al.

2010).

860 ... Eroglu

According to the literature, myxomycetes have been reported as a weak

disease on fungi growing on culture stumps (Chung & al. 1998, Lee & al.

2014). However, according to these authors, myxomycetes are not pathogenic,

rather consuming mushroom hyphae and spores as food. Although there

are studies on myxomycetes developing on cultures elsewhere (Chung &

al. 1998, Desrumaux & al. 2003), none were detected in Turkey. Although

Physarum spectabile generally commonly occurs on cacti and succulent

plants (Nannenga-Bremekamp & al. 1984), the species is reported here for

the first time on mushroom culture logs.

Acknowledgments

The author is indebted to the Scientific Research Projects (BAP) Coordinating

Office (BAP/13401030, BAP/15401116) at Selcuk University for their financial

support of the current work. I would like to thank Dr. Steven L. Stephenson

(University of Arkansas, Fayetteville U.S.A.) and Dr. Sinan Alkan (Selcuk University,

Konya, Turkey) for reviewing this article.

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htpps:// doi-org/10.5943/mycosphere/4/6/2

MY COTAXON

October-December 2021— Volume 136, pp. 865-873

https://doi.org/10.5248/136.865

First report of Podosphaera cercidiphylli

on endangered Cercidiphyllum japonicum in China

ZI-JIA PENG', SHU-YAN Liu*3, ZHONG-DONG Yu", MEI Qi™4

‘College of Forestry, Northwest A&F University, Yangling 712100, China

*Engineering Research Center of Chinese Ministry of Education for Edible and Medicinal Fungi,

Changchun 130118, China

College of Agronomy, Jilin Agricultural University, Changchun 130118, China

‘Xian LvHuan Forestry Technical Service Ltd., Xian 710018, China

* CORRESPONDENCE TO: yuzhongdong001@nwsuaf.edu.cn

ABSTRACT—A powdery mildew collected from the protected and endangered species

Cercidiphyllum japonicum was identified morphologically as Podosphaera cercidiphylli,

an identification confirmed by sequence analyses of the internal transcribed spacer (ITS)

and large subunit (LSU) of nuclear ribosomal DNA (nrDNA). This is the first report of

P. cercidiphylli in China.

Key worps—Ascomycota, Cercidiphyllaceae, Erysiphaceae, taxonomy

Introduction

Podosphaera Kunze (Erysiphaceae, Helotiales) is a large genus with 62

species and 13 varieties worldwide, including 39 species native to Europe

(Braun 1987, 1995). Thirty-six species have been reported from Japan

(Nomura 1997), 16 species from Korea (Shin 2000), and 42 species (plus five

varieties) from China (Tang & al. 2018). Within China, nine species and two

varieties in P sect. Podosphaera are known to infect 26 hosts (representing

10 genera and three families), while 33 species and three varieties in P. sect.

Sphaerotheca are pathogens of 115 different hosts (representing 65 genera in

19 families) (Tang & al. 2018).

866 ... Peng & al.

Cercidiphyllum japonicum Sieb. & Zucc. (Cercidiphyllaceae) is a plant

endemic to eastern Asia, found in only a few Asian countries including

China and Japan (Pan & Liu 1999). In China, C. japonicum is classified as an

endangered species protected for its dioecism and extremely low regeneration

rate, with natural populations distributed only sporadically in mixed forests

of the warm temperate zone and subtropical mountains (Zeng & al. 2020).

Unfortunately, plant diseases and insect pest attack threaten C. japonicum,

with additional environmental stress. Our research revealed that both

powdery mildew and mites (Eriophyidae) had jointly infected C. japonicum

trees with the chasmothecia tending to obscure patches of cell hyperplasia

caused by mites. This fungus was not easy to detect as the powdery white

symptoms were inconspicuous. When we discovered cell hyperplasia in

C. japonicum, we determined that this pathogen is new to China.

Materials & methods

Collection procedures & morphological examination

Samples of Cercidiphyllum japonicum leaves damaged by powdery mildew and

mites were collected during 2018-2020 in Ankang City, southern Shaanxi Province,

China. Herbarium specimens with chasmothecia were dried in plant presses

and deposited in the Mycological Herbarium of Forestry College, Northwest A

& F University, Yangling, Shaanxi Province, China (HMNWAFU-CF), and the

Herbarium Mycologicum Academiae Sinicae, Beijing, China (HMAS).

Chasmothecia were picked carefully from the leaves, mounted in distilled water

on a glass slide, and examined under an Olympus CX41 light microscope. Their

colour, shape, cell wall surface, appendage position, length, apex branching, septum,

shape, size, and numbers of asci and ascospores were observed and documented

in detail. We checked 50 chasmothecia taken randomly from specimens and

calculated averages and standard deviations using Microsoft Excel 2019.

Black-and-white images of the fungal teleomorph and mites were produced

with a Nova NanoSEM 450 field emission scanning electron microscope according

to Pathan & al. (2008). Partial photochromes were obtained by microphotographic

focus stacking using a Lumix DMC-G8M camera fitted with Lumix 100-300 mm

F4-5.6 67 mm and Mitutoyo M Plan Apo L 10x/0.28 0/0 f=200 lenses, M26 to

M67 adapter, Wemacro Rail’, and Helicon® Focus 7 and Zerene Stacker® software

(see https://www.wemacro.com/).

DNA extraction

Genomic DNA was extracted from the chasmothecia with BioRad Chelex-100

following published procedures (Walsh & al. 1991, Hirata & Takamatsu 1996, Bai

Podosphaera cercidiphylli on Cercidiphyllum japonicum new to China... 867

& al. 2014). Twenty-thirty chasmothecia were picked using a sterilized dissecting

needle under a dissecting microscope and mounted onto microslides. A coverslip

was used to squeeze the chasmothecia gently until broken, both slides and coverslip

were rinsed gently with 50 uL 5% Chelex-100 sterilized water solution into a 1.5

mL centrifuge tube, which was placed into a 56 °C water bath for 20 min and then

centrifuged at 5000 rpm for 1 min; the tube was then placed in a 100 °C water

bath for 8 min, vortexed vigorously, and centrifuged for 1 min at 15000 rpm. The

supernatant was transferred into another tube to be used as the template DNA.

PCR amplification

The nrDNA ITS region was amplified by nested PCR methods using ITS5 (White

& al. 1990) and P3 (Kusaba & Tsuge 1995) as the first primer pair and ITS4 (White

& al. 1990) and PM5 (Takamatsu & Kano 2001) as the second pair in 20 uL volumes

containing 10 uL 2x CoWin Taq PCR MasterMix, 1 wL each Shanghai Sangon

Biotech primer, 2 uL extracted DNA (65.9 ng/uL), and 6 uL ddH,O. The samples

were processed in a Bioer GeneAmp PCR TC-96 beginning with a 4-min initial

denaturation at 95 °C, 35 cycles of 1-min denaturation at 94 °C + 1 min at 50 °C

(1 min at 60 °C for the second pair of primers) + 1 min at 72 °C, and a final 10-min

extension at 72 °C, with the procedure ending at 4 °C.

The nrDNA large subunit (LSU)/28S region was amplified with the primers

LSU1 and LSU2 (Scholin & al. 1999) at the same volume (20 uL) and under the

same amplification conditions as the previous pair of primers ITS5 and P3. The

PCR products were separated by electrophoresis on a 1% agarose gel in TAE buffer

and purified using the Bioteke Zymoclean™ Gel DNA Recovery Kit according to

the manufacturer's instructions. The purified DNA products were amplified again

as depicted above, and the PCR products were then sequenced by Shanghai Sangon

Biotech Co., Ltd.

Phylogenetic analysis

Sequences were revised in BioEdit 7.2.5, BLAST searched in NCBI, and deposited

in GenBank (MN960597, MN960655). Other homologous sequences from the

GenBank database were downloaded for further comparisons. DNA sequences

were aligned using ClustalX vers. 1.8. (http://www-igbmc.u-strasbg.fr/BioInfo/);

gaps or missing data were eliminated from the datasets.

A neighbor-joining (NJ) tree inferred from ITS and LSU sequences was

constructed using MEGA X (https://www.megasoftware.net/home) with the

K2+G model. Maximum-parsimony (MP) and maximum-likelihood (ML)

were generated with PAUP 4.0b10 (http://phylosolutions.com/paup-test/) as

described by Jiang & al. (2011) and RAxMLGUI 2.0 (Edler & al. 2021) under a

GTR+GAMMA model. Bootstrap percentages (BP) derived from 1000 replicates

are indicated at the tree nodes.

868 ...

Peng & al.

94/96/95 Leveillula taurica AB022387

92/83/68 Phyllactinia moricola ABO22401 i Phyllactinieae

Pleochaeta turbinata AB218773

90/81/74 pete Neoerysiphe galeopsidis AB022369 |

Arthrocladiella mougeotit AB022379 Tribe Golovinomycetea

84/74/61 Golovinomyces circumfusus AB022360

S050 Erysiphe adunca var. adunca AB022374

T3168/69 Erysiphe mori AB022418 |

93/83/80 Erysiphe carpini-laxiflorae AB252472

56/-/72 | Podosphaera lini MK749431

Tribe Erysipheae

63/57/- Podosphaera spiraeae MF919434

100/100/100 52/53/59, Podosphaera pannosa MK898816

ezibulot Podosphaera aphanis MK357425

Podosphaera macularis MH687414

43/55/56] gop ¢

pe Podosphaera prunicola MG076955 Tribe Cystotheceae

Podosphaera tridactyla AB022393

86/89/93 Podosphaera xanthii AB525914

98/94/97

99/99/99 Podosphaera clandestina MG062783

69/45/39 HMAS248348 MN960597

Sawadaea tulasnei AB022366

70/64/54 Cystotheca wrightii AB022355

Blumeria graminis ABO22399 J] Tribe Blumerieae

94/87/82

-Byssoascus striatosporus AF222495 J] Out group

0.010

Fic. 1. Topological phylogram inferred from LSU rDNA from 21 taxa of Erysiphaceae with

Byssoascus striatosporus as outgroup. Bootstrap percentages from methods of neighbor-joining

(NJBPs), maximum-parsimony (MPBPs), and maximum-likelihood (MLBPs) are indicated at

nodes.

Results

LSU rDNA phylogenetic analyses

PCR amplification with LSU primers produced a unique 550 bp sequence.

The BLAST search revealed that HMAS 248348 had a high similarity to

Podosphaera spp. The phylogenetic tree (Fic. 1) clustered HMAS 248348 in

tribe Cystotheceae, separating it from the other tribes in Erysiphaceae with

strong bootstrap supports (NJBP = 99%, MPBP = 99%, MLBP = 99%). It was

closely related to P. clandestina with a 69-scored NJBP support.

ITS nrDNA phylogenetic analyses

After amplification with the ITS5/P3 primer pair, no obvious

electrophoresis products were observed due to low PCR yields. Those

amplicons served as the template for the ITS4/PM5 primer pair for nested

PCR, which produced a unique 450 bp sequence as checked with gel

electrophoresis. That sequence was aligned with 17 GenBank sequences

of tribe Cystotheceae. ITS nrDNA phylogenetic analyses (Fic. 2) clustered

HMAS 248348 and Podosphaera cercidiphylli in a monophyletic clade with

Podosphaera cercidiphylli on Cercidiphyllum japonicum new to China... 869

77/-/61 Podosphaera spiraeae AB026152

59/-/- Podosphaera filtpendulae AB022385

76/-1- Podosphaera minor AB026137

81/-/- Podosphaera pannosa AB022348

Podosphaera cercidiphylli AB026140

99/68/87 |_ HMAS248348 MN960655

85/81/88

Podosphaera leucotricha AB02723 1

Podosphaera photiniae AB026138

84/63/70 88/94/96 [-Podosphaera cayratiae ABO26151

97/96/- Podosphaera xanthit AB026146

Podosphaera intermedia AB026145

100/100/100 99/97/94 75/-/- Podosphaera elsholtziae AB026142

49/-/- Podosphaera erigerontis-canadensis AB026148

Podosphaera longiseta AB000945

Cystotheca lanestris AB000933

100/100/100 Cystotheca wrightii AB000932

Sawadaea polyfida AB000936

100/100/100 Sawadaea tulasnet AB022367

0.020

Fic. 2. Topological phylogram inferred from ITS sequences from 17 taxa of Erysiphaceae

tr. Cystotheceae with two Sawadaea taxa as outgroup. Bootstrap percentages from methods

of neighbor-joining (NJBPs), maximum-parsimony (MPBPs), and maximum-likelihood

(MLBPs) are indicated at nodes.

fairly strong bootstrap supports (NJBP = 99%, MPBP = 68%, MLBP = 87%).

Therefore, we identified HMAS 248348 as P. cercidiphylli.

Taxonomy

Podosphaera cercidiphylli Tanda & Y. Nomura,

Trans. Mycol. Soc. Japan 27: 25. 1986. Big?

Mycelia amphigenous in epidermal hyperplasia caused by mites (Eriophyes

sp.), mostly hypophyllous, evanescent or subpersistent, powdery layer not

obvious; mites aggregating on epidermal hyperplasia, massive ecdyces of

melts visible in patches; chasmothecia mostly scattered or subgregarious on

epidermal hyperplasia, rarely scattered epiphyllously, brown or dark brown,

lighter colour incipiently, globose or subglobose, 53.8-79.0 (average 67.0 +

7.4) um diam., wall cells irregularly polygonal, 6.9-18.9 x 10.4-23.8 (av. 11.4

+ 3.0 x 15.7 + 3.1) um; appendages 2-9 (mostly 3-7, av. 5 + 1.8), produced

870 ... Peng & al.

from the upper to equatorial part of the chasmothecia, radial, straight or

arcuately curved, thick-walled throughout, 1-3 times longer than the

chasmothecial diam., 70.9-226.0 (av. 129.2 + 40.8) um long, 5.3-9.3 (av. 7.1 +

0.9) um wide at the slightly thicker base, 3.3-7.2 (av. 5.3 + 0.9) um wide at the

upper part, 1-6(-7)-septate, 2-5 times dichotomously branching at the apex,

coloured dark brown from below the branch downward, ultimate branches

short and obtuse, recurved or rarely straight; ascus single, subglobose to

broadly ovate, sessile, double-walled, 41.3-62.6 x 49.6-69.1 (av. 53.5 + 4.9

x 60.7 + 4.9) um; ascospores 8, oblong or ellipsoidal, 8.9-15.4 x 14.2-24.9

(av. 12.6 + 1.9 x 19.1 + 2.7) um.

SPECIMENS EXAMINED: CHINA. SHAANXI PROVINCE: Ankang city, Zhenping

County, 32.1°N 109.2°E, alt. 1024 m, on living leaves of Cercidiphyllum japonicum, July

2018, Zhong-dong Yu (HMNWAFU-CE 2018017); July 2019, Zhong-dong Yu & Zi-jia

Peng (HMAS 248348, GenBank MN960597, MN960655); Ningshan County, 33.4 °N

108.5°E, alt. 1649 m, on living leaves of C. japonicum, September 2020, Zhong-dong

Yu & Zi-jia Peng (HMNWAFU-CEF 2020021)

DISTRIBUTION: Ankang city, Shaanxi Province, China; Hokkaido and Niigata

Prefectures, Japan.

Discussion

Cercidiphyllum japonicum is distributed in China and Japan, and

Podosphaera cercidiphylli (P. sect. Podosphaera) was originally described

from Japan (Tanda & Nomura 1986). Here, we provide the first report from

China based on morphological and phylogenetic characters; no anamorphic

forms (neither mycelial layer nor conidia) were found. Tanda & Nomura

(1986) cited 1-5-septate appendages arising from the basal region of the

perithecium, but our Chinese specimens usually developed 1-7-septate

appendages from the upper half of the equatorial region of chasmothecia,

with each appendage dichotomously branched at a right or obtuse angle,

and with each branch slightly shorter than the older one. However, Tanda

& Nomura (1986) showed appendages dichotomously developing at a right

angle with nearly an identical length for the consecutive branches. We found

that there were usually eight ascospores in a single ascus instead of seven

to eight ascospores as reported by Tanda & Nomura (1986). We believe

that these differences are attributable to variable intraspecific development,

possibly resulting from different observation methods and weather

conditions at sampling sites.

Additionally, we are the first to observe a co-infection of Cercidiphyllum

japonicum by Podosphaera cercidiphylli and mites. Chasmothecia were

Podosphaera cercidiphylli on Cercidiphyllum japonicum new to China... 871

rama f

: KS,

x360 Fee

Fic. 3. Podosphaera cercidiphylli (HMAS 248348), teleomorphic characteristics and mites

in the curled leaf: A-C. Symptoms of powdery mildew on leaf of Cercidiphyllum japonicum;

D. Mycelia and chasmothecia; E, F. Ecdyces of mites (white arrow) and chasmothecia; G. Ascus,

ascospores, and appendages; H. Insertion position of appendages; I. Dichotomous branch of

appendages; J. Mite; K. Hyperplastic epidermal cells of leaf (red arrow), mycelia (purple arrow),

mite (green arrow) and young chasmothecia (yellow arrow). Scale bars: A, B = 1 cm; C= 5 mm;

D-F = 350 um; G, H, K = 50 um; I, J = 10 pm.

mainly distributed on the abaxial surface of leaves, most of which obscure

patches of cell hyperplasia caused by mites. A Podosphaera species associated

with mites was reported by Zheng & Yu (1987) occurring on Betula pendula

Roth and B. tianschanica Rupr. infected by P. erineophila Naumov (Jaczewski

1927), but the intimate correlation between this powdery mildew and mites

is not clear. We did not find the anamorphic stage of P. cercidiphylli on our

specimens (in contrast to the Japanese specimens of Tanda & Nomura

872 ... Peng &al.

1986); we speculate that either conidia had not been produced or that mite

grazing had completely removed them. This is the first report of Podosphaera

cercidiphylli in China.

Acknowledgments

This study was funded by National Key Research Projects (2017YFD0600103-

4-2) and National Natural Science Foundation of China (31670650). We wish to

thank Prof. Susumu Takamatsu (Mie University, Japan) and Prof. Xiao-yong Liu

(Chinese Academy of Sciences, Beijing, China) for their kindness of providing

references about Podosphaera cercidiphylli. We also appreciate the expert reviews

to the early manuscript by Prof. Lu-chao Bai (Qinghai University, Xining, China)

and Prof. Xiao-yong Liu (Institute of Microbiology, Chinese Academy of Sciences,

Beijing, China).

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Sinicae 37: 1-12. (in Chinese). http://doi.org/10.11707/j.1001-7488.20010201

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plant surfaces—horses for courses. Micron 39: 1049-1061.

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Scholin CA, Marin R, Miller PE, Doucette GJ, Powell CL, Haydock P, Howard J, Ray J. 1999. DNA

probes and a receptor-binding assay for detection of Pseudo-nitzschia (Bacillariophyceae)

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35:1356-1367. https://doi.org/10.1046/j.1529-8817.1999.3561356.x

Podosphaera cercidiphylli on Cercidiphyllum japonicum new to China ... 873

Shin HD. 2000. Erysiphaceae of Korea. National Institute of Agricultural Science and Technology,

Suwon. 320 p.

Takamatsu S, Kano Y. 2001. PCR primers useful for nucleotide sequences of rDNA of the

powdery mildew fungi. Mycoscience 42: 135-139. https://doi.org/10.1007/BF02463987

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Transactions of the Mycological Society of Japan 27: 17-27.

Tang SR, Guan GX, Liu SY. 2018. Advances on the taxonomy of Erysiphales in China. Journal of

Fungal Research 16: 138-149. (in Chinese). http://doi.org/10.13341/j.jfr.2018.8006

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Zeng CY, Zhong QJ, Wang CY, Hu YP, Wu MH, Meng W, Peng MC. 2020. Ecologically suitable

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Zheng RY, Yu YN. 1987. Flora Fungorum Sinicorum, vol. 1 (Erysiphales). Science Press,

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MYCOTAXON

October-December 2021— Volume 136, p. 875

https://doi.org/10.5248/136.875

REGIONAL ANNOTATED MYCOBIOTA NEW TO THE MYCOTAXON WEBSITE:

AsstRACcT—The beautifully illustrated 30-page “Macromycetes from woodland

zones of Milpa Alta mayoralty, Mexico City, Mexico” by Sierra and eleven co-authors

may now be downloaded from MycorTaxon’s mycobiota webpage. This brings to 151

the number of free-access Fungae uploaded or linked to

http://www.mycotaxon.com/mycobiota/index.html

NORTH AMERICA

Mexico

SIGFRIDO SIERRA, SANDRA CASTRO-SANTIUSTE, IBETH RODRIGUEZ-

GUTIERREZ, ARELI E. GONZALEZ-MENDOZA, MARIO AARON GUTIERREZ-

SANCHEZ, LISETTE CHAVEZ-GARCIA, DANIELA ABIGAIL GUZMAN-

RAMIREZ, JOSE DE JESUS RUIZ-RAMOS, GUADALUPE GALVAN-BECERRIL,

NAVITH ALEJANDRA LOPEZ-GARDUZA, LILIA PEREZ-RAMIREZ, JOAQUIN

CIFUENTES. Macromycetes from woodland zones of Milpa Alta mayoralty,

Mexico City, Mexico. 27 p.

ABSTRACT— Previous studies about macromycete diversity in the Milpa Alta

mayoralty demonstrated the presence of only one species (Amanita aspera var.

franchetii). To update records from this region, specimens were collected in

woodland areas during the rainy seasons (June-October) from 2008 to 2017. A

total of 225 specimens were collected in 32 localities across different elevations

and vegetation types. Specimens were morphologically identified with

specialized literature. The macromycetes studied include 82 species, 59 genera,

and 36 families. There is one new record for the country (Calyptella campanula)

and 29 for Mexico City, and all the species are new to the mayoralty. Reference

material was deposited in the Herbarium of the Sciences Faculty, UNAM,

Mexico City (FCME). There are still unexplored localities in the area, so many

more species are expected to be discovered.

KEY worps—Ascomycota, Basidiomycota, Sierra Chichinautzin, taxonomy,

Trans-Mexican Volcanic Belt

MYCOTAXON

October-December 2021— Volume 136, p. 877

https://doi.org/10.5248/136.877

REGIONAL ANNOTATED MYCOBIOTA NEW TO THE MYCOTAXON WEBSITE:

ABSTRACT— The 10-page “A checklist of Agaricus from Pakistan” by Niazi, Ghafoor,

and Bashir may now be downloaded from Mycotaxon’s mycobiota webpage. This

brings to 152 the number of free-access Fungae available on

http://www.mycotaxon.com/mycobiota/index.html

INDIAN SUBCONTINENT

Pakistan

ABDUL REHMAN NIAZI, ANEEQA GHAFOOR, HrirRA BaAsutr. A checklist of

Agaricus from Pakistan. 10 p.

ABSTRACT—We present a comprehensive checklist of the Agaricus species

recorded from various regions of Pakistan. Thirty-two species are documented

from Pakistan according to published reports and the latest literature.

Orthographic variants and misidentified species have been excluded.

KEY worps—Agaricaceae, diversity, distribution, systematics, mycobiota

MYCOTAXON

Octoer-December 2021— Volume 136, pp. 379-380

https://doi.org/10.5248/136.379

BOOK REVIEWS AND NOTICES:

ELSE C. VELLINGA

861 Keeler Avenue, Berkeley CA 94708, USA

AsBsTRACT—We are pleased to recommend “Common edible and poisonous

mushrooms of southwestern China” by Yang, Wu, Li, Wang, and Cai. (2021).

Common edible and poisonous mushrooms of southwestern China. By

Z-L. Yang, G. Wu, Y-C. Li, X. Wang, Q. Cai, 2021. Science Press, Beijing, China.

198.00 ¥, https://life.sciencereading.cn 370 p with color plates. Hardback, in Chinese

and English. ISBN 978-7-03-068388-5.

One of the nice things to do for a mycologist traveling in

Southeast Asia is a visit to the local market to marvel at the

displays of locally collected mushrooms for sale. All those

species—various boletes, Amanita, Termitomyces, and many

others—and all in one place. Names are of course only in the

local languages, and one has to dive into the scientific literature

to find answers for one’s questions. Fortunately, there are now

several books available to help the myco-traveler getting to know the edible

and poisonous mushrooms of the area. In 2019 AN INTRODUCTION TO THE

EDIBLE, POISONOUS AND MEDICINAL FUNGI OF NORTHERN LAOs was published,

reviewed here by Vellinga (2019), and a large volume on poisonous mushrooms

for China finally saw the light in 2016 (Chen & al. 2016).

* Book reviews or books for consideration for coverage in this column should be sent to the

EDITOR-IN-CHIEF <editor@mycotaxon.com> 6720 NW Skyline, Portland OR 97229 USA.

BS: Vellinga

The book on common edible and poisonous mushrooms discussed

here focuses on one region of China only—the southwest, covering Tibet;

the provinces of Yunnan, Sichuan, and Guizhou; and the municipality of

Chongqing; this region is extremely varied, in topology, climate, and habitat.

The introduction is relatively short, only 25 pages, and focuses on the region,

the difficulties distinguishing edible species from their toxic look-alikes,

preventing and treating poisonings, descriptive terms of the mushrooms,

and the various groups and how to key those out. A key to the main groups is

included, but not one to the species.

The main section is dedicated to the species themselves, and for each species

a photo taken in the wild (a very few photos are taken outside the natural setting)

is given, complemented by a description and an end note regarding edibility or

toxicity. The species are ordered according to their general morphology (gilled

mushrooms, boletes, polypores, etc.), and edible and poisonous species are

not in separate parts. This arrangement gives the reader a better idea about

how similar poisonous species can be to edible ones. To give an idea about the

coverage, 60 bolete species and 30 amanitas are included. and Termitomyces is

represented by nine species. Nomenclature is in general up to date (although

Cystoderma aureum is still in Phaeolepiota), and the plates and descriptions are

of high quality.

A few final words. All the texts in this book are

in Chinese and in English, which makes it very

accessible and useful for those who don't read

Chinese.

tips oT }

OE ik. MEL

This book gives a very nice introduction to the | (7..%Mesnes

very rich mycoflora/funga of southwestern China,

but of course, only gives information on the edible

and poisonous species. Nonetheless, as such it is

a welcome addition to the Chinese mushroom

book library.

Chen Z-H, Yang Z-L, Bau T, Li T-H. 2016.

Poisonous mushrooms: recognition of 3

and poisoning treatment. Science Press, i

Beijing, China. 308 p.

Leessoe T, Sparre Pedersen O, Sysouphanthong P. 2019. An introduction to

the edible, poisonous and medicinal fungi of northern Laos. 192 p.

Vellinga EC. 2019. Book reviews and notices. Mycotaxon 134: 583-590.

https://doi.org.10.5248/134.583

ELSE VELLINGA

ecvellinga@comcast,net